Skip to main content

Full text of "Journal of the Lepidopterists' Society"

See other formats 


oe 
Me cea We a 
tthe 5's 


Cr 


04 Ow 


eter 
ya 


best 


25 SLOP Gy 
BH pot A 








~ 
} 
, fi 
1 yar 
i 
b 
| , 
u f 
- 











Molume 40 1986 Weather 4 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





9 October 1986 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CLIFFORD D. FERRIS, President DouGLas C. FERGUSON, 

Don R. Davis, Immediate Past President President-Elect 

JERRY A. POWELL, Vice President EDWARD M. PIKE, Vice President 
RICHARD A. ARNOLD, Secretary ALLAN WATSON, Vice President 


ERIC H. METZLER, Treasurer 


Members at large: 


JOHN M. BuRNS Boyce A. DRUMMOND III MIRNA M. CASAGRANDE 
FLOYD W. PRESTON JOHN LANE EDWARD C. KNUDSON 
JACQUELINE Y. MILLER ROBERT K. ROBBINS FREDERICK W. STEHR 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 48229, U.S.A.; and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume, is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$25.00 (institutional subscriptions) and $18.00 (active member rate) by the Lepidopter- 
ists’ Society, % Los Angeles County Museum of Natural History, 900 Exposition Boule- 
vard, Los Angeles, CA 90007. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, 
1900 John St., Manhattan Beach, CA 90266. 


Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from 
Dyar 1899, J. New York Entomol. Soc. 7:61-67. Suggested by Marc E. Epstein. 


JouRNAL OF 
Tue LeEpPIDOPTERISTS’ SOCIETY 


Volume 40 1986 Number 1] 


Journal of the Lepidopterists’ Society 
40(1), 1986, 1-7 


PRESIDENTIAL ADDRESS, 1984: 
A TRIBUTE TO THE AMATEUR 


LEE D. MILLER 


Allyn Museum of Entomology of the Florida State Museum, 
3621 Bay Shore Road, Sarasota, Florida 33580 


First of all, let me state that it has been a great pleasure to serve the 
Society as its president during the past year. This is a wonderful group 
and one of the few in this country where the professional and the 
amateur can speak as equals. But, all too often the statement is made: 
“He is only an amateur,” usually implying that he doesn’t know what 
he is talking about, or at least that his opinion is not worth as much as 
one who makes his living in the field. I do not accept this view, and 
this presentation is an unabashed tribute to those who do not make 
their living in the field of lepidopterology. 

What is an amateur? The word is derived from the Latin “amator’’ 
(lover) or the French “amare” (to love), and is defined in the dictionary 
as “one who cultivates a particular pursuit, study or science from taste, 
without pursuing it professionally.”” Everyone makes his or her living 
at something, and I consider an amateur lepidopterist as one who makes 
his living at something other than lepidopterology. There can, then, 
be janitors, pipefitters, medical doctors, engineers and mammalian 
ecologists using lepidopterology as an avocation. There are good am- 
ateur workers in the field and bad ones, but the variant definition of 
amateur is not always applicable: “‘one practicing an art without mas- 
tery of its essentials.’ I am praising that amateur who has gained a 
certain amount (often a great deal) of expertise in a particular phase 
of lepidopterology, enough so that he or she is able to impart that 
knowledge to others. 

Amateurs have been the backbone of science since its inception: 
there were no professional entomologists in Linnaeus’ time, but no one 
suggests that they did not do the best possible job with the information 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


available to them. Pieter Cramer was an artist whose avocation was 
lepidopterology; Dru Drury was a silversmith; Jacob Huebner was a 
printer; William Chapman Hewitson was a wealthy landowner and 
otherwise retired; William Henry Edwards was a coal baron; Henry 
Edwards was an actor; William Barnes was a physician, and Walter 
Rothschild was a “‘black sheep” who did not fit into the family banking 
business. From these diverse backgrounds, however, came people who 
made tremendous contributions to the field of lepidopterology, as great 
as those of the contemporaneous professionals. Most of the contribu- 
tions that have been made by amateurs have been in systematics, be- 
havior, life history, morphology and ecology, subsets of the field that 
usually require less elaborate equipment; but a number of important 
contributions also have been made in the field of genetics by non- 
professionals. 

Often the amateur will attack a problem of limited scope and be- 
come very expert on that topic. The amateur may be expected to 
gather much pertinent material within a circumscribed group, often 
more and better material than is readily available to the professional 
even in the finest of facilities. He studies the material gathered, in- 
cluding that which is borrowed from museum collections, and gathers 
together the appropriate literature, just as the professional would. He 
is at least as likely as the professional to ask opinions from others, and 
most studies done by amateurs go through a great many more drafts, 
and hands for comment, than do papers written by those of us in the 
profession for the simple reason that the amateur is still convinced that 
he does not know everything there is to know. Most publishing ama- 
teurs are at least as receptive to new and different ideas as are those 
of us who do it for a living. 

Admittedly, I am talking here of the best amateurs, not those un- 
guidable persons who nevertheless write on various subjects without 
the benefit of knowledge or guidance. The “good’’ amateurs are the 
ones who behave in the field about as we should expect professionals 
to perform their studies. 

The early amateurs worked chiefly in systematics of Lepidoptera. 
They generally amassed huge (for the day), usually beautifully curated 
collections of specimens, and spent much of their time writing descrip- 
tions of new taxa from these collections or from those of their ac- 
quaintances. Certainly this is what W. C. Hewitson, W. H. Edwards 
and Henry Edwards, to name a few, did, and their works compare 
favorably to those of Butler, Walker and other professionals of the day. 
Once in a while one hears grumblings about the incompleteness by 
today’s standards of their descriptions, but one also hears this complaint 
about the descriptions done by professionals of the same time period. 


VOLUME 40, NUMBER 1 3 


Later, other amateurs like Frederick DuCane Godman and Osbert 
Salvin, two wealthy English “men of leisure’, began looking at Lepi- 
doptera as related populations of organisms, rather than as specimens 
which either varied from other specimens or did not. These amateurs 
began putting contemporary biological theories into practice and crit- 
ically examined lepidopteran populations in light of the then new ideas 
of evolution and biogeography. The result was that they, mostly with 
Godman’s money, decided to do a total biological inventory of Mexico 
and Central America for which they would write some parts and enlist 
experts in other fields to contribute sections. The resulting Biologia 
Centrali-Americana was published over more than 30 years and ran 
to nearly fifty sumptuous volumes. This publication still has not been 
superceded. | 

Walter Rothschild, once he decided to collect Lepidoptera seriously, 
could have been expected to do it with a vengeance. He did, finally 
amassing something over two and a quarter million specimens, more 
than the Lepidoptera holdings of most of today’s major museums. He 
immediately saw the advantages of collecting study series, and for the 
first time, a private collection had more than a few specimens of any 
single taxon. At the same time in France, a printer, Charles Oberthuer, 
began much the same type of accumulation of material, although the 
scope of his collecting was smaller than Rothschild’s. Oberthuer’s col- 
lection finally amounted to more than a million specimens. Rothschild 
was one of the first private collectors to realize that he probably needed 
professional curatorial help, and he hired Karl Jordan as entomologist 
at his museum and Ernst Hartert as his ornithological curator. What a 
team they made! If one goes through the writings of this triumvirate, 
one can find the first modern concept of geographical subspecies, tri- 
nominal nomenclature, an elucidation of the biological species concept, 
and one of the first cladistic analyses of an animal group (it was not so 
labelled and frequently is not recognized as such). Especially with 
Walter Rothschild and his curators, modern systematics can be shown 
to have had its birth. To say that I am a Rothschild fan would be true 
because of the facts that he (A) collected long series of material for 
study, (B) had a worldwide bias to his activities, and (C) surrounded 
himself with those who in conjunction with him developed most of the 
bases of modern systematics. Their efforts were not always appreciated 
by their contemporaries, but those must have been halcyon days at 
Tring! 

Strangely enough, another man lived in England at the same time 
who had an obsession with outdoing Rothschild, thereby setting himself 
a prodigious task. James J. Joicey was a man of leisure who decided 
that he would outdo Rothschild in the acquisition of orchids. He tried 





4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


this for some years before and during World War I, finally going 
bankrupt for 30,000 pounds, a very large sum at the time. The judge 
admonished Joicey, and he agreed that he would not try to build the 
world’s premier orchid collection, and he held to his promise. He 
switched instead to Lepidoptera and went broke in the 1930's for over 
300,000 pounds, perhaps making a statement about how deeply one 
can get involved in making a collection if one lets his or her imagi- 
nation run wild. Joicey did, however, hire curators (perhaps because 
Rothschild had them), and they produced some excellent work, espe- 
cially on the Lepidoptera of New Guinea, Hainan Island, and central 
and eastern Africa. The Joicey, Oberthuer and Rothschild collections 
are the reason that the British Museum (Natural History) enjoys such 
numerical superiority over other collections throughout the world. 

In the United States, Dr. William Barnes, a physician from Decatur, 
Illinois, formed a magnificent collection of North American Lepidop- 
tera. It became readily apparent that many specimens that he was 
obtaining were undescribed, and with the help of professional curators, 
he set about to describe them. From this activity came revisionary 
studies on moth genera which are still standard today. Many of these 
were published in his own journal, Contributions to the natural history 
of the Lepidoptera of North America, and were often jointly coau- 
thored with his curators, Arthur Ward Lindsey, James McDunnough, 
and Foster H. Benjamin, among others. Many of the finest contribu- 
tions during the first quarter of the 20th Century came from that 
journal. 

In later years, many amateurs have made contributions to the tax- 
onomy of Lepidoptera. Without exception, these students have been 
willing to listen to others and be guided by them, and the resulting 
papers have been highly informative. Anyone who has ever studied 
Hesperiidae knows of the legacy left us by Brigadier William Harry 
Evans, a retired British army officer who served in India during the 
first third of this century. His Catalogues to the hesperiids are standard 
works for the professional and the amateur alike. Many people forget 
that he also wrote the definitive guide to the butterflies of India, which 
is still being reprinted. Still, he was an amateur who performed like a 
professional. Another military man turned lepidopterist is John N. Eliot, 
recently awarded the Karl Jordan Medal, whose studies of the Lycaeni- 
dae and the Neptis group of nymphalids have earned him a permanent 
place in our field. He also completely revised Corbet and Pendlebury’s 
Butterflies of the Malay Peninsula. Our former President, Col. Stanley 
S. Nicolay, has done some fine work tying together certain of the 
neotropical hairstreaks, and he is still trying to bring some order out 
of the chaos that characterizes this group. Stan also works on the hes- 


VOLUME 40, NUMBER 1 5 


periids, so it might be said that he has selected some very knotty 
problems to tackle. Surely the taxonomy of North American moths is 
better because of the activities of Mr. André Blanchard, a retired en- 
gineer. I have the utmost admiration for one who can tackle these little 
creatures and make sense of them. The late M. Henri Stempffer was a 
retired French banker who has added greatly to our knowledge of the 
taxonomy of African Lycaenidae, a group that has puzzled all workers 
before him. For his contributions M. Stempffer was elected to receive 
the first Jordan Medal given by this Society. We can expect other 
amateurs to receive the award in the future, I am sure. Arthur Rydon 
studied the Charaxini and has increased our knowledge of this group 
through his writings. Similarly, the recently deceased Dr. Lionel G. 
Higgins, an English physician, made a lifetime study of the Melitaei- 
nae, and his writings are the basis of the classification of this group 
throughout the world. Our own former President and Honorary Life 
Member, Dr. F. M. Brown, is a geologist by trade, but he has achieved 
renown in the taxonomy of both nearctic and neotropical Rhopalocera, 
and if this is not enough, Brownie has now engaged (at more than 80 
years old) in the study of fossil insects, and is writing several compendia 
on important groups. Cliff Ferris, another amateur subsequently elect- 
ed President of our Society, is a bioengineer by profession, but his 
studies on the systematics of Nearctic butterflies are quite professional. 
Let us not forget the contributions of a former lawyer, Dr. Cyril F. 
dos Passos, who has worked for several decades on the taxonomy and 
nomenclature of North American butterflies. The list of amateurs who 
have contributed to the taxonomy of Lepidoptera is endless, and I 
apologize to others that I have left out—there simply is not enough 
space. 

A special place must be saved for the late Dick Dominick whose 
dream of The Moths of North America has been an inspiration to us 
all. The several volumes that have appeared under the aegis of this 
series conspire to make the study of moths as popular as that of but- 
terflies. It was an impressive project, and one can but wish that Dick 
had lived to see more of it completed. 

To finally get to nontaxonomic studies, the life history studies done 
by Roy and Connie Kendall, their associates, and by those such as Dave 
Baggett have added the biological information that can turn alpha 
taxonomic treatises into studies at a higher level. They are providing 
the building blocks for greater understanding of Lepidoptera, and these 
studies cannot help but improve the quality of later studies on these 
insects. Amateur life history work generally does not include the chae- 
totaxy of the larvae that professionals consider vital, but the workers 
mentioned here are also preserving egg, larval, and pupal material for 


6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


such studies, and this material is readily available to those interested 
in performing them. 

Dr. David Wright is a physician by occupation, still he is embarking 
on some superb micromorphological studies of various stages of but- 
terflies, especially the Lycaenidae, using scanning electron microscopy, 
and he has already published new and innovative ideas on the mor- 
phology of the larval stages, including showing that some setae have 
wrongly been attributed to some segments when they belong to adja- 
cent ones. To the casual observer, these data may seem trivial, but he 
has found at least one derived (apomorphic) character which seems to 
define the Lycaeninae, one that was previously unsuspected. 

There are other examples of amateurs making contributions in ecol- 
ogy, physiology, and even genetics of Lepidoptera. The information 
that these workers are providing surely aids in the understanding of 
these insects. An example of such a study is Mike Fisher’s rearing of 
Papilio nitra which unexpectedly turned up the fact that Papilio zel- 
icaon gothica is the more frequent color morph of nitra. This was an 
elegant study, done on a low budget, that showed something totally 
unexpected. 

My wife and I have been fortunate enough to work for and with a 
man with vision much like that of Rothschild. The late Arthur Allyn 
was the heart of an effort to form a real resource for taxonomic and 
morphological research on Macrolepidoptera. In the years since we 
joined him, that collection grew from about 100,000 specimens to its 
present roughly 550,000 prepared and 250,000 unspread specimens. 
The decision was made to gather material from throughout the world 
because we felt that revisionary studies done on groups from only part 
of their ranges were incomplete, and we wished to encourage a world 
view of butterflies and large moths. Another goal was to obtain collec- 
tions in their entirety and not see them broken up as so many had been 
in the past. The vision of Art Allyn has been an inspiration to us, and 
we hope that the material he was responsible for gathering will be used 
for generations to come. 

Thus far, almost everyone that I have mentioned in this tribute has 
been male. Consider the fact that the Rothschilds are and were an 
amazing lot, true “Renaissance men’. The present Renaissance “man” 
of the Rothschilds is a woman: Miriam Rothschild has published a 
quarter of a million words on flea taxonomy, she has worked on plant- 
insect interactions, on intestinal worms, on mimicry in butterflies, on 
the behavior of sea gulls and has been involved with worldwide con- 
servation efforts. Miriam Rothschild was educated basically at home, 
chiefly by her uncle Walter and his curators at Tring. She earned no 
university degree, but nonetheless is honored in scientific circles in 


VOLUME 40, NUMBER 1 7 


England and elsewhere. Her ability and tenacity have made her 
achievements possible. 

The honor role of amateurs is long and distinguished. They have 
made and are making significant contributions to our chosen field. The 
next time you hear the statement, ““He’s only an amateur,” realize that 
this should not be a pejorative; perhaps it is a tribute, since the person 
in question does not have to be paid to perform. All of us, amateurs 
and professionals alike, have something to give lepidopterology. The 
difference between amateur and professional is one of degree, rather 
than kind. In the final analysis, there is not “amateur science” and 
“professional science’, there is only good science or poor science. Let 
us recognize that we all have something worthwhile to say, and we 
will all benefit from such understanding. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 7 


GENERAL NOTE 


TRYON REAKIRT: A SEQUEL 


In 1964 I wrote briefly about Tryon Reakirt, a Philadelphia entomologist of note during 
the 1860’s (Brown 1964, J. Lepid. Soc. 18:211-214). He was a mystery man in his last 
years. All I knew earlier was that he had fled the country in early 1871. As a result, both 
his enterprises and his father’s business filed for bankruptcy. I found the answer to his 
disappearance among newspaper clippings belonging to William Henry Edwards of 
Coalburgh, West Virginia. 

One clipping is from the Philadelphia Inquirer, Wednesday, 8 February 1871. Reakirt 
had forged notes on large pharmaceutical houses to the tune of more than $110,000! An 
error in a date caused a bank clerk to go into the matter with the purported issuer. The 
fat was in the fire! Reakirt left town hurriedly, and ultimately got to Lima, Peru, where, 
apparently, he died of dysentery in late 1872 or early 1873. 


F. MARTIN BROWN, 6715 South Marksheffel Road, Colorado Springs, Colorado 80911. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 8-19 


A NEW SPECIES OF EPIDROMIA (NOCTUIDAE) 
FROM FLORIDA 


M. ALMA SOLIS 


Maryland Center for Systematic Entomology, Dept. of Entomology, 
University of Maryland, College Park, Maryland 20742 


ABSTRACT. This preliminary study of the genus Epidromia Guenée (Ophiderinae: 
Noctuidae: Lepidoptera) describes a new species E. fergusoni and redescribes the type 
species E. pannosa Guenée. 


Twenty species group names are included in the neotropical genus 
Epidromia Guenée (Ophiderinae). This genus is distributed in north- 
ern South America and throughout Central America and reaches its 
greatest diversity in the Antilles. Florida is its northernmost extension. 


METHODS 


Names applied to wing veins and markings correspond to Forbes 
(1923). The forewing length is measured from the base of the wing to 
the apex of the wing. The width is measured from the apex to the anal 
angle. The depth and width of the cleft of the ostium bursae is mea- 
sured as shown in Fig. 1. All measurements correspond to the mean 
value, and the measurements in the parentheses are the range. The 
numbers in parentheses in the distribution section are USNM (United 
States National Museum of Natural History) genitalia slide numbers. 


RESULTS 


Epidromia is being redescribed since the original description is in- 
complete. The genitalia are described and illustrated for the first time. 


Epidromia 


Epidromia Guenée, 1852, In Boisduval and Guenée, Hist. Nat. Insectes, Spec. Gén. des 
Lépid. 7:325. Type: Epidromia pannosa Guenée, 1852, by subsequent designation 
(Berio, 1966. Annali Museo Civico Storia Naturale Giacomo Doria 76:57). 


Penultimate segment of palpi upturned and longer than first and third segments to- 
gether; ultimate segment ending in blunt point. Abdomen cylindrical, elongated; sternites 
more hairy than tergites; distal end appearing square-shaped in males and tapered in 
females (Figs. 3, 6 and 8) distal tergite square-shaped with membranous projections into 
seventh tergite, projections pointing medially; distal sternite with two sclerotized, long 
lobes connected by two smaller lobes (Fig. 2). Front legs stout with dense hairs. Wings 
entire, oblong; underside of wings having beige sheen with silky pubescence; peak of 
each scallop of adterminal line with immediately adjacent dot. Sacculus simple; uncus 
simple, widely angled distally; tegumen medially extended to point ventro-laterally; 
vinculum about as long as wide; vesica distally bilobed without cornuti (Figs. 4, 5 and 
9, 10). Genital plate heart-shaped; ostium bursae cleft; no signa on corpus bursae (Figs. 
7 and 12). 


VOLUME 40, NUMBER 1 9 





Fic. 1. Diagrammatic illustration of ostium bursa and measured distances. 


Inspection of specimens and representative genitalia slides of genera 
believed to be closely related revealed that Epidromia is most closely 
related to Itomia Htibner. These two genera share an uncus that is 
simple, curved, and with a prominent spine at the distal end, and a 
costa on the valve that is strongly lobed. Itomia species more closely 
resemble Epidromia species from South America than those from Cen- 
tral America and the Antilles. 

The type-species of Itomia is Itomia lignaris Hiibner, by monotypy. 


1.0 mm 





Fic. 2. Last abdominal segment of Epidromia. 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 3. Epidromia fergusoni, holotype, male. 


The holotype is believed to have been destroyed (R. W. Poole, pers. 
comm.). The original description and illustration of I. lignaris are not 
adequate for identification of the species. The specimens at the USNM 
suggest that a complex of species is involved; therefore, a male and a 
female from the complex have been chosen for the purpose of the 
genitalia comparison (Female: Mizantlan, Mexico, slide no. 42,618; 
Male: Mizantlan, Mexico, slide no. 42,619) (Table 1). 

Table 2 is simply a list of species group names currently placed in 
Epidromia. Some names are new combinations. Some of the names 
have been traditionally recognized as synonyms. Some species de- 
scribed by Walker from the Antilles are suspected to be synonyms; he 
described only females, and females are highly variable. Photographs 
of the Walker and Gueneé types from the British Museum were stud- 
ied. There are three other new species that will not be described here. 
A full revisionary treatment is not feasible at this time because of the 
difficulties of assembling sufficient material and identifying all of the 


types. 


Epidromia fergusoni Solis, new species 


Male holotype. Scales of collar, thorax and abdomen brown with purplish tinge (pur- 
plish tinge may disappear with age). Third sternite of abdomen without large bristles 
(n= 7). Ground color of wings brown with purplish tinge. Forewing: Other margin 
undulated, invaginated at end of R;, forming a peak at apex. Basal band brown (not 
visible); antemedial line light brown; triangular, chocolate brown area extending distally 
from base of antemedial line, terminating where postmedial line curves toward base of 
wing. Reniform spot gray (black, more obvious in older specimens). Postmedial line beige; 


VOLUME 40, NUMBER 1 


TABLE 1. 


Structure 


Ultimate labial palp 
segment 

Penultimate labial 
palp segment 

Tergites and ster- 
nites of abdomen 

Distal end of abdo- 
men 


Front legs 
Underside of wings 


Dots at peak of 
each scallop of 
adterminal line 

Upper side of wings 

Sacculus 


Tegumen 
Vinculum 
Uncus 
Vesica 


Genital plate 


Ostium bursae 


Epidromia 


blunt 


four times as long as ultimate 
palp segment 

sternites with more hair than 
tergites 

distal tergite square with 
membranous projections 
into 7th tergite bending 
dorsally; distal sternite with 
medial sclerotized area but 
not tubelike 

stout, with dense long hairs 

beige sheen with silky pubes- 
cence 

immediately adjacent 


variable 
simple 


medially extended to a point 
ventrolaterally 

as long as wide 

not sharply angled 


distally bilobed 
heart-shaped 


with a definite cleft 


11 


Comparison of Epidromia and Itomia. 


Itomia 


pointed 


two times as long as ultimate 
palp segment 

sternites and tergites equally 
hairy 

distal tergite triangular with 
membranous projections into 
7th tergite bending laterally; 
distal sternite with medial 
tubelike sclerotized area 


slender, devoid of long hairs 

yellow without silky pubes- 
cence 

not immediately adjacent 


diagonal lines across the wings 

chitinized extension costad and 
extending distally 

simple 


longer than wide 
sharply angled distally 


not distally bilobed 


rectangular, slightly longer 
than wide 
cleft not definite 


chocolate brown patch adjacent to postmedial line beginning at M, and extending distally 
to the apex of forewing. Adterminal line light brown, peak of each scallop with beige 
dot. Terminal line light brown (beige). Distal underside of wing without gold patch 
between M, and R,. Forewing length 2.3 cm (2.2—2.4) (n = 25). Length/width ratio 1.5 
(1.8-1.7). Hindwing: Outer margin round, area adjacent to margin light brown. Post- 
medial line same as in forewing; chocolate brown area extending from postmedial line 
to about halfway to base of wing (Fig. 3). Genitalia: Uncus enlarged at distal end; valve 
with thumblike process on saccular margin, distal end of valve expands into small, flaplike 
process; editum on costa round; longest lobe of vesica bifurcate, with short branch round- 
ed and longer branch rounded; longer branch expanded at base and tapered to blunt 
point (n = 7) (Figs. 4, 5). 

Female allotype. Scales of collar, thorax and abdomen brown. Ground color of wings 
brown. Forewing: Outer margin same as in male. Basal line (if visible) double, dark 
brown on inside, yellow on inside; median line dark brown (or absent). Reniform spot 
same as in male. Postmedial line double, brown on inside and yellow outside. Amount 
of beige in area between postmedial and adterminal line when present varies. Subter- 
minal line yellow to brown with dot at peak of each scallop, terminal line brown. Fore- 
wing length 2.1 cm (1.9-2.2) (n = 25). Length/width ratio 1.5 (1.4-1.6). Hindwing: 
Outer margin same as in male. Line markings same as forewing, but basal line and 
antemedial line not visible (Fig. 6). Genitalia: Ostium bursae cleft, approximately 0.45 
mm wide at distal end, depth of cleft 0.85 mm (0.28-0.40) (n = 4) (Fig. 7). 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1.0 mm 


1.0 mm 





5 


Fics. 4 & 5. Male genitalia of Epidromia fergusoni. 4, Egmont, Fla., USNM slide 
42,611; 5, aedeagus, Miami, Fla., USNM slide 42,609. 


Types. Holotype: Male (Fig. 3), University Reserve, Welaka, Putnam Co., Florida, 6 
April 1972, D. C. Ferguson. Allotype: Female (Fig. 6), University Reserve, Welaka, 
Putnam Co., Florida, 6 April 1972, D. C. Ferguson. Paratypes: Six males, same locality 
and collector, USNM genitalia slide numbers 42,054, 42,614, 42,607; three females, same 
locality and collector. All specimens are deposited in the U.S. National Museum of Nat- 
ural History, Washington, D.C. 

Distribution. Other specimens used in this analysis were from the following localities 
in Florida: Miami, Glenwood, Ft. Myers, Ft. Meade, Dade City, St. Petersburg, Royal 
Palm State Park, Marcos Island, Lutz, Stemper, Indian River, Egmont, Ft. Lauderdale, 
Chokoloskee. Specimens with suspect data: Plainsfield, N.Y.; Jemez Springs, N.M.; Cuba. 


VOLUME 40, NUMBER 1 13 





Fic. 6. Epidromia fergusoni, allotype, female. 


Discussion 


This is the Florida species that has been known in this country for 


nearly a century as Epidromia delinquens (Walker) (=Ophiusa delin- 
quens Walker, 1858). However, Hampson (1913) referred delinquens 
to the synonymy of Mocis repanda (F.), a name that Hampson and 


TABLE 2. List of species group names in Epidromia. (Names in parentheses are the 


original combinations. ) 


Epidromia pannosa Guenée 


POSS 


. zetophora Guenée 

. xanthogramma Wallengren 
. zephyritis Schaus 

. rotundata Herrich-Schaffer 
. consperata Dognin 


poaphiloides Guenée 

profana Walker 

flavilineata (Hampson) NEW COMBINATION (Thermesia flavilineata) 
glaucescens (Walker) NEW COMBINATION (Thermesia glaucescens) 


. lenis (Walker) NEW COMBINATION (Thermesia lenis) 
. antica (Walker) NEW COMBINATION (Ophisma antica) 


arenosa (Walker) NEW COMBINATION (Phurys arenosa) 


. pedestris (Walker) NEW COMBINATION (Phurys pedestris) 

. profecta (Walker) NEW COMBINATION (Poaphila profecta) 

. saturatior (Walker) NEW COMBINATION (Remigia saturatior) 

. sigillata (Walker) NEW COMBINATION (Thermesia sigillata) 

. suffusa (Walker) NEW COMBINATION (Thermesia suffusa) 

. tinctifera (Walker) NEW COMBINATION (Thermesia tinctifera) 
. valida (Walker) NEW COMBINATION (Ophisma valida) 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1.0 mm 





Fic. 7. Female genitalia of Epidromia fergusoni, paratype, Welaka, Fla., USNM 
slide 42,615. 


other authors subsequently (but mistakenly) considered to represent 
the same species as Mocis latipes (Guenée). The type of delinquens 
has not been seen, but after reading the description by Walker, it is 
acknowledged that Hampson was correct in assigning it to the genus 
Mocis Hubner. 


It should be noted that E. fergusoni is not the only species of Epi- 


VOLUME 40, NUMBER 1 15 





Fic. 8. Epidromia pannosa Guenée, holotype, male. 


dromia in Florida. A male specimen from Homestead of one unde- 
scribed species and a male and a female from Big Pine Key of another 
undescribed species have also been collected there and are now re- 
corded from the continental United States for the first time. 

E. fergusoni is named after Douglas C. Ferguson who collected the 
type material. 


Epidromia pannosa Guenée 


Epidromia pannosa Guenée, 1852, In Boisduval and Guenée, Hist. Nat. Insectes, Spec. 
Gén. des Lépid. 7:326. 


TABLE 8. Comparison of E. pannosa and E. fergusoni. 


Overali color 
Third sternite 


Structure pannosa 


brown 
with bristles 


fergusoni 


purplish 
without bristles 


Outer forewing margin straight undulated 

Antemedial and postmedial line double single 

Triangular, chocolate brown area extending absent present 
distally from base of antemedial line 

Hindwing outer margin—male angulate rounded 

Hindwing outer margin—female rounded rounded 

Thumblike process on costa absent present 

Short branch of bifurcated lobe of vesica truncated rounded 

Cleft of ostium bursae 0.62 mm wide, 0.25 mm wide, 

0.43 mm depth 0.35 mm depth 
On underside of forewing—gold patch be- present absent 


tween M, and R, 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1.0 mm 





1.0 mm 





10 


Fics. 9 & 10. Male genitalia of Epidromia pannosa Guenée. 9, Rio Janeiro, Brazil, 
USNM slide 42,053; 10, aedeagus, Venezuela, USNM slide 42,065. 


Male. Scales of thorax and abdomen brown; head and collar dark brown. Third sternite 
with bristles (n = 4). Ground color of wings brown. Forewing: Outer margin not undu- 
lated. Basal band brown; antemedial line double, yellow on inside (may not be evident) 
and dark brown on outside. Median shade dark brown. Reniform spot outlined in dark 
brown, filled with gray (black or gray without outline). Postmedial line double, dark 
brown inside, yellow outside. Subterminal line faint, light brown (chocolate brown or 
lacking). Dark brown patch between subterminal and adterminal line extending from 


VOLUME 40, NUMBER 1 17 





Fic. 11. Epidromia pannosa Guenée, female. 


M, to anal angle. Adterminal line dark brown, slightly scalloped; peak of each scallop 
with dark brown dot. Terminal line yellow (brown). (One specimen with vein lines dark 
brown.) Distal underside of wing with gold patch between M, and R,. Forewing length: 
2.3 cm (2.1-2.4). Length/width ratio: 1.6 (1.4-1.8) (n = 10). Hindwing: Outer margin 
slightly angulate at end of Cu,; edge of wing above Cu, parallel to body. Area from apex 
to anal angle adjacent to outer margin dark brown (light brown or no shading). Median 
shade, postmedial and subterminal line same as forewing (Fig. 8). Genitalia: Uncus 
enlarged at distal end; editum on costa round; vesica with longest lobe bifurcates, short 
branch truncate, long branch expanded at base, tapering to blunt point (n = 4) (Figs. 9 
and 10). 

Female. Since Gueneé did not describe the female of this species, the following de- 
scription of a female believed to represent the same species from Aroa, Venezuela is 
provided. Three males of this species were collected at Aroa, Venezuela and all four 
(three males and one female) have the same label information. Scales of head, collar, 
thorax, and abdomen same color as male. Ground color of wings same as male. Forewing: 
Median line dark brown. Reniform spot same as male. Postmedial line same as male; 
subterminal line beige. Length: 2.1 cm. Length/width ratio: 1.7 (n = 1). Hindwing: 
Outer margin round. Median line, postmedial line and subterminal line same as forewing 
(Fig. 11). Genitalia: Ostium bursa cleft, approximately 0.62 mm wide at distal end, depth 
of cleft 0.48 mm (n = 1) (Fig. 12). 

Types. The holotype, in the British Museum of Natural History, is a male with no 
label data other than “‘Bresil.’”” A photograph of the type specimen taken by R. W. Poole 
was used in this description. 

Distribution. Aroa, Venezuela: one female (42,115) and three males (42,065); Edo. 
Zuela, Venezuela: three males; Rio Janeiro: one male (42,053); Castro, Parana: one male 
(42,057); Merida, Mexico: one male (42,069); Tamazunchale, Mexico: four males; Ma- 
zatlan, Mexico: one male (42,056); Poza Rica, Mexico (42,072). All specimens examined 
are in the collection of the U.S. National Museum of Natural History. 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





1.0 mm 


eee 
Q 
te 
Sey 


2 


Fic. 12. Female genitalia of Epidromia pannosa Guenée, Venezuela, USNM slide 
42,115. 


Discussion 


J. G. Franclemont and E. Todd (1983) mistakenly synonymized pan- 
nosa and poaphiloides. This synonymy was probably based on notes 
by Hampson that were never published and transferred from the Brit- 
ish Museum of Natural History to the U.S. National Museum of Natural 
History. E. poaphiloides was described from Cayenne, French Guiana 
by Gueneé in 1852. The type of poaphiloides is believed to have been 
destroyed or lost (R. W. Poole, pers. comm.). An illustration of the type 


VOLUME 40, NUMBER 1 19 


is not available. After reading the written description, it is quite ob- 
vious that poaphiloides is not pannosa. A number of specimens of 
poaphiloides from French Guiana and British Guiana (Guyana) can 
be found at the U.S. National Museum of Natural History. 


ACKNOWLEDGMENTS 


I wish to thank Dr. R. W. Poole, SEL, USDA, for the use of the catalogue of the 
Noctuidae at the U.S. National Museum of Natural History, for photographing the types 
at the British Museum of Natural History and for instructing me in dissection techniques. 
I thank Dr. D. Miller and Dr. D. Ferguson, SEL, USDA for corrections and suggestions 
on the manuscript. This description was prepared for a course taught by Dr. D. Miller 
at the University of Maryland at College Park. Mr. A. M. Wilson photographed the 
specimens. I thank Mrs. Elaine R. S. Hodges, Smithsonian Institution, for instructing me 
in illustration techniques and for analyzing my illustrations. Scientific Article No. A- 
4064, Contribution No. 7049 of the Maryland Agricultural Experiment Station. 


LITERATURE CITED 


BERIO, E. 1966. Nomi Generici polispecifici di Noctuidae del globo con scelte di specie 
tipo e observazioni. Annali del Museo Civico di Storia Naturale “Giacomio Doria” 
76:57. 

FORBES, W. T. M. 1928. Lepidoptera of New York and neighboring states. Part I. 
Cornell Univ. Agri. Expt. Sta. Mem. 68:19-30. 

FRANCLEMONT, J. G. & E. L. Topp. Noctuidae. In Hodges, R. W., et al. 1983. Check 
list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd. and The 
Wedge Entomological Research Foundation, London. xxiv + 284 pp. 

GUENEE, A. 1852. Noctuelites III. In Boisduval et Guenée, Histoire naturelle des in- 
sectes, species géneral des Lépidoptéres. 7:215-216, 325-326. Librairie Ency. de 
Roret, Paris. 

Hampson, H. G. 1913. Catalogue of the Noctuidae in the collection of the British 
Museum 13:84. Board of Trustees. London. 

HUBNER, J. 1823. Zutrage zur Sammlung Exotischer Schmettlinge, (etc.), 2. Augsburg. 
32 pp. + 8 numbered pages, figures 201-400, 33 plates (1819-1822). 

KIMBALL, C. P. 1965. The Lepidoptera of Florida. Arthropods of Florida and neigh- 
boring land areas, I: v + 363 pages, 26 pl. 

NyE, I. W. B. 1975. The generic names of the moths of the World. Vol. 1: Noctuidae, 
Agaristidae, Nolidae. Trustees of the British Museum, London. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 20-22 


THE LARVA AND PUPA OF LYCOREA PIETERI 
LAMAS (DANAIDAE) 


DAVID KENNETH WETHERBEE 


San José 71, Restauracién, Republica Dominicana 


ABSTRACT. The larva and pupa of the danaid Lycorea pietri Lamas is described 
and figured for the first time. It differs most markedly from the larvae of other danaids 
in having only a pair of tentacles in front and none in the rear. The food plant is Carica 
papaya (Cariaceae). 


During the last quarter of the 18th Century there was a gifted ar- 
chitect in northern Haiti who was also a naturalist 200 years ahead of 
his time. His work is unknown to biologists even though he painted 
from life accurately hundreds of Haitian animals and plants. Most of 
his subjects did not become known to binomial taxonomy until two 
generations later. He is unpublished, except that apparently Deshayes 
pirated some of his work and sent it to Buffon (Wetherbee, in press). 
Not only did he paint some 50 Lepidoptera, but he reared them from 
larvae and depicted the early stages and named several of them from 
their food plants. 

He was ““M. de Rabié, marechal de camp, ingenieur en chef de la 
parties du nord de St. Domingue” and resided at Cap Haitian from at 
least as early as 1752 to about 1784 and died in Paris in 1785. His first 
insects were drawn in 1766. Folios of his work are now in the Blacker- 
Wood Library of McGill University. Through the courtesy of Miss 
Eleanor MacLean, McGill Librarian, I have been privileged to publish 
Rabié’s zoological subjects (Wetherbee, 1985a). 

Rabié reared Lycorea pieteri Lamas (formerly called L. ceres Cra- 
mer, 1779) and depicted the larva, pupa, shed “‘robe’’ and imago. He 
called the larva ‘“‘chenille de papayer (food plant: Carica papaya of 
the Caricaceae) and the adult “le noeud de ruban’”’ (ribbon-bow). This 
is L. pieteri cleobaea Godart, 1819, the type-specimen of which was 
collected, undoubtedly, by Antonio Gonzales of the covert Baudin voy- 
age to Hispaniola in 1799. 

Most of the larvae of Lepidoptera drawn by Rabié were those of 
moths, especially of Sphingidae (Wetherbee, 1985b). Only four other 
butterfly species of the 36 species illustrated by Rabié (Wetherbee, 
1985a) show early stages: Danaus plexippus, Colobura dirce, Siproeta 
stelenes and Dione vanillae. 

As can be seen from Fig. 1 (the black and white reproduction hardly 
does justice to the beauty of Rabié’s colored painting), the larva (per- 
haps fourth instar) is danaid in character, but unlike Danaus, which 
has pairs of both anterior and posterior fleshy tentacles, and unlike 


VOLUME 40, NUMBER 1 x1 


C 11 N ata ) . ° a“ 


\ 





Le Noeunpr Ruspan. 





Fic. 1. Lycorea pieteri Lamas (L. p. cleobaea Godart, 1819) life history as painted 
by Rabié in Haiti in 1782. 


Anetia which has none, Lycorea pieteri has only a pair of anterior 
tentacles. These are slightly longer than those of Danaus plexippus. 
The larva is smooth, about the same size as D. plexippus; both the 
head and posterior segment are black; the thoracic segments are white, 
followed by seven golden-yellow segments and then one white next to 
the last segment in front of the black tail-end. The narrow black bands, 
one at the anterior of each segment, have short, black, single, lateral, 
oblique, dash-like marks pointing backwards and downwards. 

The hanging pupa is shown by Rabié in its lateral aspect only. It is 
similar in texture, size and shape to D. plexippus (but without the 
ridge) and is golden yellow with black bump-dots running in two arched 
lines on the sides, a few on the anterior parts, one prominent “‘occipi- 
tal” bump-dot and two anal ones. The cremaster is black and contrasts 
sharply with the white web. 

One must pause to consider that this excellent work was accom- 
plished contemporary with Linnaeus’ 12th edition of the Systema Na- 
turae in a country which has had essentially no entomological research 
up to the present time. If we consider that Audubon was somewhat of 
a pioneer and hero, certainly Rabié was even a greater one. 

Since viewing Rabié’s pictures, I have had the good fortune in No- 


22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


vember of finding many of the larvae of this species in Restauracion, 
Republica Dominicana feeding on Carica papaya. The early stages are 
only slightly tinged with yellow. The fifth instar does not have the 
posterior segments white as shown by Rabie, but the thoracic segments 
tend to be whitish. The black tentacles are 15 mm in length, and the 
larva is about 40 mm. The ten narrow, black bands on the anterior of 
each segment send a spur toward each black spiracle and sometimes 
include it. The ultimate yellow segment has only three black dots 
representing the band. There are a pair of yellow eye-spots on the 
small black ultimate black segment. Both in Rabié’s illustration and in 
life, it is easy to mistake the tail for the head in the resting caterpillar. 

The pupa is always suspended from the midvein of the green lechosa 
leaf about midway along the length of the leaf. It is dull waxen-yellow. 
The black dots run in a mid-dorsal series, a dorso-lateral series, and 
there is an arc of elongated dots on the wing following the curvature 
of the wing and an isolated black dot in the middle of the wing. As 
shown by Rabié, there is a pair of black bumps on the body near the 
base of the cremaster. 


LITERATURE CITED 


WETHERBEE, D. K. 1985a. Zoological exploration of Haiti for endemic species. Pub- 
lished privately, Shelburne, Massachusetts. 556 pp. 

1985b. The sphinx-moths (Sphingidae) of Hispaniola and the 18th century moth 

paintings of Rabié. Published privately, Shelburne, Massachusetts. 69 pp. 





Journal of the Lepidopterists’ Society 
40(1), 1986, 23-26 


LIFE HISTORY AND HABITS OF EXOTELEIA ANOMALA 
HODGES, A PONDEROSA PINE NEEDLE MINER IN THE 
SOUTHWESTERN UNITED STATES (GELECHIIDAE) 


ROBERT E. STEVENS! 


Rocky Mountain Forest and Range Experiment Station, 
USDA Forest Service, Fort Collins, Colorado 80526 


ABSTRACT. Exoteleia anomala, the larvae of which mine needles of ponderosa pine 
in Arizona and New Mexico, has a one-year life cycle similar to that of several species 
of needle-mining Coleotechnites. Each larva requires two needles to complete devel- 
opment. When numerous, larvae can cause highly visible foliage damage, but outbreaks 
do not appear to persist. 


In summer 1977, I reared an undescribed species of Exoteleia (Ge- 
lechiidae) from foliage of ponderosa pines, Pinus ponderosa Dougl. ex 
Laws., near Silver City, Grant Co., New Mexico. In 1978, entomologists 
with the USDA Forest Service, Southwestern Region, Albuquerque, 
New Mexico, reported a needle miner infestation in ponderosa pines 
from an area near Show Low and Pinetop, Navajo Co., Arizona, some 
90 km NW of Silver City. No moths were obtained at that time, but 
a resurgence of the population in 1981 provided material for study; it 
too was E. anomala. Collections and observations in 1981 and 1982, 
reported here, have made it possible to outline the species’ life history 


and habits. 


METHODS 


Collections of foliage representing at least two years’ growth were 
made 21 October 1981, and 4 March, 6 April, 10 May, 22 June, 12 
July, and 25 August 1982 for the 1981-82 generation, and a single 
collection from the 1982-83 generation was obtained on 11 November 
1982. Life history events and larval habits were recorded following 
examination of the foliage, and sufficient numbers of larvae were pre- 
served to permit determination of instars. Adult voucher specimens are 
deposited in the U.S. National Museum of Natural History. 


DESCRIPTION 


The adult is described in detail by Hodges (1985). Briefly, it is a 
small, fragile moth, forewing length 4-5 mm, with a whitish head, 
mottled gray-brown to black and white forewings, and mottled brown 
and whitish abdomen. The hindwings are grayish, and have fringes of 
long hairs. 


‘Present address: Dept. of Entomology, Colorado State University, Ft. Collins, Colorado 80523. 


24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


First-stage larvae are tan, with light brown head capsules and pro- 
thoracic and anal shields. Instars 2, 3, and 4 are brown, with dark 
brown to black head capsules and anal shields. Prepupal fourth-stage 
larvae are dark brown to almost black. 

Its appearance, coupled with its needle-mining habit, can lead 
nonspecialists to confuse this species with well known pine needle min- 
ers in the genus Coleotechnites. 


LIFE HISTORY AND HABITS 


Exoteleia anomala has a one-year life cycle. The moths fly and 
oviposit in midsummer, and the larvae overwinter. Pupation takes place 
in the mined needles. The life history is similar to that of a more 
northern but potentially sympatric species of needle miner Coleo- 
technites ponderosae Hodges & Stevens (Stevens 1973, Hodges & Ste- 
vens 1978), and an undescribed species of Coleotechnites from Pinus 
jeffreyi Grev. & Balf. in southern California (Luck 1976). However, 
larval habits of E. anomala are different. 

Adults fly in June and July. Eggs were not seen, but they may be 
laid in old mined needles as with Coleotechnites pine needle miners 
(Stark 1954, Struble 1972, Stevens 1978), or in other locations near 
susceptible new foliage. 

Larvae readily colonize foliage of the current year’s growth, in con- 
trast to C. ponderosae larvae, which seem to prefer older foliage (Ste- 
vens 1973). Examination of shoots from 12 heavily infested trees col- 
lected 21 October 1981 showed that 72% of the new (1981) needles 
had been invaded, as had 65% of the 1980 foliage. How much of the 
damage to 1980 foliage resulted from mining by the 1981 generation 
was not determined; certainly some was attributable to the 1980 gen- 
eration. A collection of 14 1981 shoots, made in July 1982 after all 
larval feeding had ended, showed that only 8% of the needles had 
escaped infestation. 

Larval head capsule measurements indicate four instars (Fig. 1). 
Each larva utilizes two needles to complete development. First instars 
enter the first needle in late summer, molt, and remain there as second 
instars until spring. Most of the larvae in the 4 March 1982 collection 
appeared to have only recently arrived at the second needle. Larvae 
enter the first-mined needles in the middle third of the needle. This 
mine is short, only 1-2 cm; the part of the needle distad of the mine 
soon dies and fades. Larvae normally enter the second-mined needle 
within 1 cm of the tip, and more of the second needle is excavated. A 
set of 28 fully developed mines (from which adults had emerged) 
averaged 5.7 cm long (SD = 1.2 cm). The larvae cut small holes in the 
needle surface for disposal of frass; these are covered with silk from 


VOLUME 40, NUMBER 1 95 


oO 


l 
aks) YV/ Y 
/ h 
/, —A Wy 
"| Y; 1; YY 
rw 6 
yy i Wi, ae 
HHUA om VA VAL 


£2428 732 ce ‘444 748. 52° 56 
Head capsule widths (mm) 


Fic. 1. Head capsule measurements (n = 138) of Exoteleia anomala larvae. 


within after they are no longer needed. There is usually only one 
needle miner per needle, but more than one larva per needle does 
occur. When this happens, normal entry locations are altered. Larvae 
located distally to others in a single needle do not complete develop- 
ment, and may move to another needle. 

Pupation, as in the well known Coleotechnites pine needle miners 
and some species of Exoteleia, takes place within the last mined needle. 
The larva cuts a hole in the needle surface to allow for adult exit. The 
hole may be at either end of the mine. Pupae are dark brown to black, 
cylindrical, and 5.5 to 6.0 mm long. They are usually found 1 cm or 
more back from the exit hole, head pointing toward it. 


POPULATION FLUCTUATIONS AND EFFECTS OF 
LARVAL FEEDING 


Although heavy larval feeding can cause many needles to die, no 
permanent tree damage has been reported. This may be due to the 


26 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


fact that outbreaks of E. anomala appear not to persist. For example, 
population densities were high enough to attract attention in Arizona 
in 1978, but no infestation was reported during the next 2 years. Larval 
numbers in 1981 appeared to decline markedly at the time of transfer 
from first- to second-mined needles. This may be a time in the insect’s 
life cycle when it is particularly vulnerable. 


ACKNOWLEDGMENTS 


I thank J. M. Schmid and J. C. Mitchell for making foliage collections, several of my 
co-workers for helpful comments on the manuscript, and R. W. Hodges for describing 
the species. 


LITERATURE CITED 


HopcEs, R. W. 1985. A new species of Exoteleia reared from ponderosa pine (Lepi- 
doptera: Gelechiidae). J. Lepid. Soc. 39:139-144. 

HonpcEs, R. W. & R. E. STEVENS. 1978. Two new pine-feeding species of Coleotechni- 
tes (Gelechiidae). J. Lepid. Soc. 32:118-122. 

Luck, R. F. 1976. Bionomics and parasites of a needle miner, Coleotechnites sp., 
infesting Jeffrey pine in Southern California. Env. Entomol. 5:937—942. 

STARK, R. W. 1954. Distribution and life history of the lodgepole needle miner (Re- 
curvaria sp.) (Lepidoptera: Gelechiidae) in Canadian Rocky Mountain Parks. Can. 
Entomol. 86:]-12. 

STEVENS, R. E. 1973. A ponderosa pine needle miner in the Colorado Front Range. 
USDA For. Serv. Res. Note RM-228, 3 pp. Rocky Mt. For. and Range Exp. Stn., 
Fort Collins, Colo. 

STRUBLE, G. R. 1972. Biology, ecology, and control of the lodgepole needle miner. U.S. 
Dep. Agric. Tech. Bull. 1458, 38 pp. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 27-35 


BIOLOGY AND IMMATURE STAGES OF HEMILEUCA DIANA 
AND H. GROTEI (SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge #111D, Houston, Texas 77054 


ABSTRACT. The primary host plant of Hemileuca diana in Arizona is Quercus 
oblongifolia, Mexican blue oak. Adult flight records extend from August to late Novem- 
ber, but peak emergence is in October. The primary host plant of H. grotei in central 
Texas is Quercus fusiformis, live oak. Adult grotei fly from late October to December. 
Adult Hemileuca grotei from New Mexico are similar to those from Texas. Immature 
stages and adults of both species are illustrated, as is the holotype of H. diana. Although 
closely related, hybrid matings between these two species do not produce viable ova. 


Hemileuca diana Packard 


Hemileuca diana is locally abundant in the mountains of Arizona 
but frequently difficult to locate or capture. Presently diana is known 
from Arizona, New Mexico, Colorado, and Sonora, Mexico. Although 
there are two old Texas records, the data are incomplete and probably 
in error (Ferguson 1971). Hemileuca diana is associated with the mon- 
tane oak habitat above 1,100 m (Fig. 1b) and is undoubtedly wide- 
spread from northern Mexico to Colorado. Because of its similarity to 
H. grotei Grote & Robinson, the two species have been confused in 
the literature, making it difficult to accurately determine the extent of 
either species distribution. 

The specimens of diana illustrated by Ferguson (1971) are typical 
in appearance but slightly smaller than average. The forewing length 
of males from southern Arizona ranges from 23 to 28 mm, X = 24.3 
mm (N = 34); females from 27.6 to 31.5 mm, X = 29.9 mm (N = 14). 
The fore- and hindwing ground color of the female is black or dark 
brown. The forewing of the male varies from brown to dark brown, 
while the hindwing is dark brown. Both sexes have a cream-colored 
medial line which passes distal to the forewing discal spot (Figs. 2a, b, 
i, j). The genitalia of a male diana was illustrated by Ferguson (1971) 
but was accompanied with grotei locality data. Ferguson (pers. comm.) 
re-examined the specimen and confirmed its identity as diana. 

Minor geographical variation has been observed among the males in 
southern Arizona. This might be expected since many of the mountain 
ranges diana inhabits are isolated by distance and habitat. Males from 
the Huachuca Mts. in Santa Cruz and Cochise counties exhibit the 
most contrast between fore- and hindwing coloration (Fig. 2a). Speci- 
mens from the Santa Catalina Mts., Pima Co. (Fig. 2b), and Graham 
Mts., Graham Co., have forewings usually, but not always, darker than 
those from the Huachuca Mts. Specimens from the Chiricahua Mts., 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. la-e. Immature stages of Hemileuca diana. a, mature larvae, lateral and dorsal 
view; b, H. diana habitat, Pima Co., Arizona; ec, egg ring on Quercus oblongifolia; d, 
ventral and lateral view of male pupa; e, pupal chamber. 


Cochise Co., are slightly smaller and have the least contrast between 
fore- and hindwings. Even though subtle differences are noted, over- 
lapping phenotypic variation among these populations makes the ap- 
plication of subspecific names unwarranted. 

Adults from central Arizona (Figs. 2c, d) appear intermediate to 
diana and grotei. The cream colored medial forewing line is thin and 
disrupted by the discal spot, as in grotei (Figs. 2e, f, g), but the hind- 


VOLUME 40, NUMBER 1] 29 








f 


; ) ; 
Se Arua atk, 


Pham Cucih, Lac 











Fics. 2a-m. Adults of Hemileuca diana, H. grotei, and immature stages. a, H. diana 
6, Huachuca Mts., Cochise Co., Az; b, H. diana 4, Santa Catalina Mts., Pima Co., Az; e, 
Hemileuca sp. 6, Oak Creek Canyon, Coconino Co., Az; d, Hemileuca sp.? 4, Sunflower, 
Maricopa Co., Az; e—h, H. grotei 66, Burnet Co., Tx; i, Holotype °, H. diana; j, H. diana 
9, Santa Catalina Mts., Pima Co., Az; k, H. grotei 2, Burnet Co., Tx; 1, mature H. grotei 
larvae, lateral view; m, mature H. groiei larvae, dorsal view (larvae from Burnet Co., 
ix), 


wing medial line is well developed, and continues to the anal wing 
margin, as in diana (Figs. 2a, b). The forewing length of the six males 
examined ranged from 21 to 24 mm. The genitalia are variable but 
have distinct characteristics. I reared one larva collected on scrub oak 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in Oak Creek Canyon, Coconino Co., Arizona. The mature larva was 
grayish, the intersegmental area was brown, and appeared distinct 
when compared to diana larvae from southern Arizona, and more 
similar to grotei larvae from Texas. Females from this population have 
not been available for examination. The material from central Arizona 
appears distinct and may represent an undescribed taxon. 

An examination of the diana holotype from Plum Creek, Colorado 
(Fig. 2i) confirms that there is no obvious difference between the type 
and females from southern Arizona (Fig. 2j). Ferguson also noted that 
the type agreed well with material from southern Arizona. Therefore, 
if a new name is proposed it should be applied to the central Arizona 
phenotype. 


Biology 


In southern Arizona, the flight period of diana extends from mid- 
September to late November, with the peak near mid- to late October, 
but this may vary from one mountain range to the next by as much as 
three weeks. Adults emerge in the morning and usually mate before 
1200 h. Male flight activity usually occurs between 0930 and 1630 h. 
Females oviposit during the afternoon. The flight of the female is 
slower, more straight, and higher among the oaks than that of the male, 
which is rapid, erratic, and usually within 3 m of the ground. 

Females deposited 80 to 140 ova in two to four separate egg rings 
near the tips of the branch close to the leaf clusters (Fig. lc). Field 
collected egg rings contained 25 to 65 greenish-gray ova. The eggs 
overwinter, and larvae usually hatch during April or early May. 

The larval hostplant in Pima, Santa Cruz, Graham and Cochise coun- 
ties, is Mexican blue oak, Quercus oblongifolia Torr. Michael J. Smith 
(pers. comm.) indicates that larvae are occasionally found on Emory 
oak, Q. emoryi Torr. Early instar larvae are black and feed gregari- 
ously. During the first two or three instars, larvae prefer developing 
flower buds and new leaves. If disturbed fourth instar larvae release 
their grip, fall to the ground, and disperse. 

Mature larvae feed singly and do not drop to the ground when 
disturbed. Some larvae have six rather than five instars but are identical 
in appearance to those with one less. The mature larva is gray with a 
distinctly dark gray dorsal area, and purple intersegmental areas (Figs. 
la, b). Larvae from the Santa Catalina and Huachuca Mts. were reared 
on five different occasions and mature larvae from the Graham Mts. 
were examined. Within these three populations, the larval phenotype 
is uniform. Pupation usually occurs in early June but James S. Mc- 
Elfresh (pers. comm.) has found mature larvae during early September. 


VOLUME 40, NUMBER 1 | 


Larvae pupate under leaf litter, where they construct a small chamber 
of debris tied together with silk (Fig. le). 

On four different occasions a total of seven diana females from 
Arizona attracted and mated with wild male grotei in Texas. Collec- 
tively, the diana females deposited nearly 900 ova, but none hatched. 
Dissection of the hybrid ova about one month after pure diana and 
grotei ova hatched revealed that only a few contained dead, partially 
developed embryos; most ova appeared to be infertile. The high degree 
of genetic incompatibility between these two taxa leaves no doubt that 
they are distinct species. Females of both H. juno (Packard) and H. 
electra (Hwy. Edwds.) can be used to attract H. diana males (Tuskes 
1984). A cross between an electra female and a male diana produced 
fertile ova, but upon hatching the larvae refused to feed on host plants 
of either species. Females of electra will also attract and mate with 
male H. eglanterina (Boisduval) but only infertile ova are produced 
(Collins & Tuskes 1979). 


Larval Description 


The larval description is based on 26 larvae reared to maturity from 
ova collected in 1982 by Mike Smith and the author, at Molino Basin, 
Santa Catalina Mts., Pima Co., Arizona. Preserved larvae are in the 
author’s collection. 


First instar. Head: Black, diameter 0.7 mm. Body: Length 6 mm, width 1.4 mm. 
Ground color black. Dorsal area black, lateral and ventral surfaces dark brown to black. 
Dorsal and dorsolateral scoli forked near apex, one seta on each fork. All scoli black. 
True legs and prolegs black. 

Second instar. Head: Black, diameter 1.5 mm. Body: Length 10-11 mm, width 2.0- 
2.2 mm. Similar to first instar except for a small red dot between the dorsolateral and 
lateral scoli on abdominal (A) segments Al and A7. 

Third instar. Head: Black, diameter 2.1-2.3 mm. Body: Length 19-20 mm, width 
4.0-4.2 mm. Ground color black. All scoli black and branched. Lateral surface with 
traces of 3 incomplete lines extending length of larva. Line 1 incomplete, undulating, 
and white, touching lateral scoli and extending length of larva. Line 2 thin, white dash 
just anterior of dorsolateral scoli on each abdominal segment. Line 3 thin, white, and 
broken by black segmental area, passing just ventral to dorsal scoli. Spiracles, prolegs, 
and true legs black. 

Fourth instar. Head: Black with short white secondary setae; diameter 3.5-3.9 mm. 
Body: Length 32-37 mm, width 9.5-11 mm. Ground color black. Three broken white 
lateral lines extend length of larva. Line 1 passes between lateral scoli and is well defined 
only on posterior portion of each segment. Line 2 passes between dorsolateral scoli; 
prominent on posterior of each segment. Line 8 passes mid-way between dorsolateral 
and dorsal scoli; broken by intersegmental area. Area between line 1 and ventral surface 
gray; between lines 1 and 2 (spiracular area) black; between lines 2 and 3, dark gray. 
Dorsal and mid-dorsal area black. All scoli black with white or hyaline colored spines. 
Short white secondary setae extend from white or light gray pinacula on lateral and 
dorsal surfaces. Prominent red-orange dot occurs between lateral and dorsolateral scoli 
on Al; similar but smaller dot on A7. Spiracles black. All shields black. Inner portion of 
proleg and ventral surface brown. True legs black. 


32 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fifth instar. Head: Dark brown with numerous white secondary setae; diameter 5.1- 
5.4 mm. Body: Ground color dark gray. Length 47-58 mm, width 7.5-10 mm. Dorso- 
lateral, lateral and sublateral scoli with black shafts and black and gray spines; base of 
shaft ringed with gray. Dorsal metathoracic scoli to A8 rosette type with black tips and 
dark gray base. Pro- and mesothoracic dorsal scoli enlarged. Two prominent light gray 
lateral lines extend length of larva and divide dorsal and lateral areas. Line 1 undulating, 
and passes between lateral scoli. Line 2 passes just ventral to dorsolateral scoli and is 
interrupted by maroon intersegmental area. Lateral segmental area between lines 1 and 
2 gray with numerous light gray pinacula which may contain short white secondary 
setae. Dorsal area black with fewer light gray pinacula than lateral segmental area. 
Intersegmental area between lateral and dorsolateral scoli black or dark gray. Dorsal 
intersegmental area maroon. Ventral surface light brown to flesh with light gray pinacula 
in intersegmental area. Prominent shiny orange spot occurs posterior and near to spiracles 
on Al and A7. Thoracic shields black. True legs dark brown. Prolegs gray with dark 
gray shields. Spiracles orange. 


Hemileuca grotei Grote & Robinson 


Until recently, little information was available on the biology of 
Hemileuca grotei. Ferguson (1971) illustrated both grotei and diana, 
and pointed out major differences in adult morphology, wing pattern, 
and distribution. Kendall and Peigler (1981) provided additional in- 
formation on the distribution and flight period of grotei in Texas. No 
mention has been made of the extreme phenotypic variation found in 
the adults, and only a partial description of the immature stages has 
been published. 

Most males and females have a well defined white medial band on 
the forewing interrupted by the dark discal spot. On the male hind- 
wing, the white medial line is usually widest between M1 and M2, 
then narrows and terminates between Cul and 2A (Figs. 2e, f, g). In 
females, the medial hindwing line is more developed, seldom strongly 
tapering, and extends beyond 2A to the anal margin of the wing (Fig. 
2k). 

Some adults are almost entirely black, with only the white bar in 
the center of the forewing discal spot present. Others have only a trace 
of the hindwing medial line (Fig. 2h). Of approximately 400 males 
examined, 3 to 7% represent the dark phenotype. The occurrence of a 
dark phenotype is not uncommon in many Hemileuca species (Tuskes 
1984). Normally, the ground color of the male is dark brown to nearly 
black. Sometimes the base of the forewing is dark gray, and the medial 
and distal portions are light gray (Fig. 2g). The ground color of the 
female is dark brown to black. Forewing length of males from Inks 
Lake, Burnet Co.; Texas, ranged from 22.0 to 25.3 mm, X = 24.2 mm 
(N = 30); females varied from 26.0 to 29.4 mm, X = 28.1 mm (N = 
30). 

In addition to Texas, Hemileuca grotei occurs in New Mexico. Rich- 
ard Holland collected a series at Dome Lookout (Sandoval Co., X-11- 


VOLUME 40, NUMBER 1 33 


84, elev. 2,460 m), and in the northwestern corner of the state at Fort 
Windgate (McKinley Co., IX-30-1975). The average forewing length 
of the Dome Lookout males is identical to that of central Texas pop- 
ulations. There are subtle differences in coloration and the frequency 
of various phenotypes between central Texas and New Mexico material 
examined. In 38% of New Mexico males (N = 18), the white medial 
hindwing line is absent; thus the wing is solid black. Further, there 
does not appear to be a relationship between the presence or absence 
of the medial hindwing line and the development of the forewing 
medial line. In Texas populations, if the hindwing line is reduced or 
absent, the forewing line also tends to be reduced. One specimen ex- 
hibits the same grayish scaling on the forewing as illustrated in Figure 
2g. The only female examined was identical to those from central 
Texas. ; 


Biology 


Both males and females are active day flyers. The flight season in 
central Texas extends from October to early December, but peak emer- 
gence is near mid- to late November. Local climatic conditions signif- 
icantly influence duration and extent of daily adult flight. In late No- 
vember 1982, 26 males were collected during a one-day trip. The high 
temperature for the day was 14.4 C, with light rain, strong gusty winds, 
and 85% cloud cover. The first males were observed at 1020 and the 
last at 1480 h. During a trip in 1988, conditions at the same location 
were clear with a high of 25 C and light winds. At that time males 
were very abundant, and in flight from 0910 to 1800 h. Females were 
observed and captured in flight from about 1200 to 1800 h. 

Newly emerged larvae are black and feed gregariously. The dark 
coloration may aid thermoregulation and increase activity during the 
early spring. Like those of H. diana, fourth instar grotei larvae tend 
to drop from the branch if disturbed. During the late fourth and fifth 
instars, larvae exhibited almost equal preference for flowers or leaves. 
The natural larval hostplant is Quercus fusiformis Small. Kendall and 
Peigler (1981) reported that Q. havardii Rydberg x Q. stellata Wan- 
genheim, Q. texana Buckley, and Q. marilandica Muenchhausen are 
also utilized, but to a lesser extent. Mature larvae measure 39 to 48 
mm in length, and have a gray ground color. The dorsal surface is 
darker than the lateral surface and the intersegmental area is reddish 
brown (Fig. 2], m). 

Before pupation the larva appears to darken, shrink in size, and the 
intersegmental color becomes less prominent and the light yellow pi- 
nacula become light gray. Larvae wander from one to three days be- 
fore constructing a loosely woven cocoon in the leaf litter. Seventy- 


34 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


seven larvae were reared from ova to maturity; 43 were males and 34 
were females. All appeared to have five larval instars, and exhibited 
little variation. As with H. diana, most grotei pupae produce adults 
the same year, but both species have the ability to spend two years in 
the pupal stage. 


Larval Description 


The description is based on 77 larvae reared to maturity from ova 
deposited by a female collected by the author at Inks Lake State Park, 
Burnet Co., Texas. 


First instar. Head: Black, diameter 0.8 mm. Body: Length 6 mm, width 1.2 mm. 
Ground color black to dark brown. Dorsal pro- and mesothoracic scoli forked at tip. 
Metathoracic scoli forked or spiked, remaining scoli spikelike. All scoli black with light 
colored setae extending from apex of each shaft. Ventral surface dark brown to black. 
True legs and prolegs black. 

Second instar. Head: Black, diameter 1.4 mm. Body: Length 10 mm, width 1.8 mm. 
Similar to first instar with one exception: A small orange patch occurs between the 
dorsolateral and lateral scoli of Al. 

Third instar. Head: Black, diameter 1.8-2 mm. Body: Length 18-19 mm, width 3.4 
mm. Ground color black. Pro- and metathoracic scoli enlarged; shaft black, spines yellow 
and black. Dorsal abdominal scoli black; spines yellow with black tips, rosette pattern 
developing. Dorsolateral, lateral and sublateral scoli black with black spines. Trace of 3 
incomplete light yellow lines extend length of larva, lines variable. Line 1 undulating, 
subspiracular, extending from Al to AQ, and touching base of each lateral scolus. Line 2 
broken, consisting of dots passing along base of dorsolateral scoli. Line 3 often well 
developed, extending from T2 to A8, passing just ventral of dorsal scoli, often disrupted 
by intersegmental area. Orange dot on Al and smaller dot on A7, both set between 
dorsolateral and lateral scoli. True legs, prolegs, and spiracles black. Small yellow pinacula 
occur on all segmental areas, including ventral surface. 

Fourth instar. Head: Black, diameter 3.0-3.4 mm. Body: Length 31-33 mm, width 
5 mm. Ground color black. Dorsal thoracic scoli T1 and T2 elongated, with black shafts 
and gold spines. Dorsal abdominal scoli rosette type with black base and gold spines, and 
light brown to black tips. Dorsolateral, lateral and sublateral scoli with black shaft and 
gold spines. Three incomplete light yellow lines extend length of larva, some may be 
poorly developed. Line 1 undulating, subspiracular, connecting base of all abdominal 
lateral scoli. Line 2 lightly marked, sometimes absent, in line with dorsolateral scoli on 
segmental area. Line 8 passes just ventral to base of dorsal scoli; interrupted by black 
intersegmental area. Body covered with small light yellow pinacula with a short hyaline 
setae extending from each. Mid-ventral surface dull orange-brown. Spiracles orange. 
Prominent orange dot occurs between lateral and dorsolateral scoli of Al; smaller dot 
similar in color and location on A7. True legs and prolegs black. 

Fifth instar. Head: Black with short white secondary setae, diameter 4.6-5.0 mm. 
Body: Length 39-48 mm, width 7.0-8.5 mm. Ground color gray. Dorsal prothoracic 
scoli (T1) elongated; shaft black with black and light yellow to cream colored spines. 
Dorsal metathoracic scoli similar to dorsal T1 scoli but with light yellow to yellow-gray 
rosette spines at base. Dorsal abdominal and TS8 scoli rosette type with yellow to yellow- 
gray spines and black tips. Dorsolateral, lateral, and sublateral scoli with black shaft and 
light yellow to white spines. Two to three incomplete light yellow to light gray lateral 
lines extend length of larva; the two dorsalmost may be poorly developed. Line 1 well 
developed, undulating, subspiracular, extending from A2 to A9, and touching base of 
each lateral scolus. Line 2 lightly marked, sometimes absent, in line with dorsolateral 
scoli on segmental area. Line 3 passes just ventral to base of dorsal scoli; interrupted by 
intersegmental area. Lateral surfaces gray, dorsal area dark grayish-black. Body covered 


VOLUME 40, NUMBER 1 35 


with small light yellow to cream or light gray pinacula with a short hyaline seta extending 
from each. Lateral intersegmental are brown to rust. Spiracles light orange. Prominent 
brown dot occurs between lateral and dorsolateral scoli on Al and A7. Ventral surface 
light brown. Thoracic shield black. True legs and prolegs dark brown to near black. 


Kendall and Peigler (1981) published a partial description of a ma- 
ture grotei larva but did not give the source of their material. Com- 
parison of their larval description with larvae from Burnet Co. indi- 
cates a number of differences. Larvae from Burnet Co. have a shiny 
black head; reddish brown intersegmental area; scoli of three different 
configurations and size; and brownish orange spiracles. Kendall and 
Peigler described grotei larvae as having a rusty brown head with 
mottled black patches; maroon intersegmental area; almost equally 
developed scoli; cream colored spiracles; and concluded they were 
most similar to larvae of H. burnsi (Watson). The larval description of 
burnsi by Comstock (1937), together with my observations suggest 
there is little similarity between these two species. In coloration and 
morphology, groeti larvae from Burnet Co., Texas are most similar to 
diana and diana-like larvae from central Arizona. 


ACKNOWLEDGMENTS 


I thank James S. McElfresh, Michael J. Smith, Kenneth C. Hansen, Richard Holland, 
Douglas Ferguson, and Donald E. Bowman for providing data or material. I also thank 
Mike Collins and Ann Tuskes for their comments and suggestions on the manuscript. 


LITERATURE CITED 


COLLINS, M. M. & P. M. TusKEs. 1979. Reproductive isolation in sympatric species of 
day flying moths (Hemileuca: Saturniidae). Evolution 33:728-733. 

ComsTOCK, J. A. & C. M. DAMMERS. 1937. Notes on the early stages of three California 
moths. Bull. So. Calif. Acad. Sci. 36:68-78. 

FERGUSON, D. C. 1971. The Moths of America north of Mexico. Fascicle 20.2a Bom- 
bycoidea (in part). Classey, London, pp. 1-154. 

KENDALL, R. C. & R. S. PEIGLER. 1981. Hemileuca grotei (Saturniidae): Natural his- 
tory, spatial and temporal distribution. J. Lepid. Soc. 35:41-50. 

TusKEs, P. M. 1984. The biology and distribution of California Hemileucinae (Satur- 
niidae). J. Lepid. Soc. 38:281-309. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 36-53 


NATURAL HISTORY AND ECOLOGICAL CHEMISTRY OF 
THE NEOTROPICAL BUTTERFLY 
PAPILIO ANCHISIADES (PAPILIONIDAE) 


ALLEN M. YOUNG 


Invertebrate Zoology Section, Milwaukee Public Museum, 
Milwaukee, Wisconsin 53233 


MURRAY S. BLUM 
Department of Entomology, University of Georgia, Athens, Georgia 30602 


HENRY M. FALES AND Z. BIAN 


Laboratory of Chemistry, National Heart, Lung and Blood Institute, 
Bethesda, Maryland 20014 


ABSTRACT. The life cycle, behavior, and chemical ecology of the neotropical but- 
terfly Papilio anchisiades idaeus Fabricius (Papilioninae) were studied, using larvae from 
a single cluster of eggs obtained in NE Costa Rica. The butterfly places large clusters of 
eggs on the ventral surface of older (bluish green) leaves of Citrus. The larvae are 
cryptically colored and exhibit communal resting, molting, and nocturnal feeding be- 
havior. Fourth and fifth instars perch on branches and trunk of the host plant. Larvae 
are parasitized by the braconid wasp Meteorus sp., and the ant Camponotus rectan- 
gularis attacks and kills pupae located on the host plant. Paper wasps do not attack large 
larvae or pupae, even though their nests are often abundant in Citrus trees occupied by 
P. anchisiades. Pupae on substrates away from the host plant may survive ant predation. 
Larvae readily evert the osmeterium when provoked; a very pungent, disagreeable odor 
is noticeable (to humans) only in the fifth instar. The principle components of the os- 
meterial secretions change both qualitatively and quantitatively with the molt to the 
fifth instar. The major corstituents of the secretions of third and fourth instars are 
sesquiterpenes including “a-bergamotene’’, a-acoradiene, “‘a-himachalene’’, and isomers 
of farnesene; the main secretion of the fifth instar is dominated by isobutyric acid and 
2-methylbutyric acid with sesquiterpenes, aliphatic hydrocarbons, long-chain alcohols, 
and carboxylic esters constituting minor constituents. The possible adaptive significance 
of this shift in the chemistry of the osmeterial defensive secretion is discussed. 


The neotropical butterfly Papilio anchisiades idaeus Fabricius (Pa- 
pilionidae: Papilioninae) is well known in Mexico, Central and South 
America (Seitz 1908, Ross 1964a, b). It is a large tailless swallowtail 
with velvety-black wings bearing white patches dorsally on the fore- 
wings, and deep red to lavender blotches dorsally on the hindwings. 
This butterfly is commonly seen around clumps or groves of Citrus 
trees (Rutaceae), the host plant of the caterpillars (Stoll 1781, Carac- 
ciolo 1981, Dewitz 1878, Moss 1919). The life cycle and early stages 
have been incompletely described (Caracciolo 1891, Dewitz 1878, Ehr- 
lich & Ehrlich 1961, Jones 1881, Moss 1919, Oliveira 1977, Ross 1964a, 
Stoll 1781). In this paper we describe and illustrate the early stages, 
and present new information on the behavior of immature stages, on 
parasitism, predation, and egg placement. In addition, we analyzed 
osmeterial secretions of third, fourth and fifth instar larvae to compare 


VOLUME 40, NUMBER 1 37 


the chemistry of these defensive secretions with those of other species 
of Papilio. Since recent investigations demonstrated both qualitative 
and quantitative changes in the secretions between Papilio fourth and 
fifth instars of other species (Seligman & Doy 1972, Burger et al. 1978, 
Honda 1980a, b, 1981), we wondered if this was also the case for P. 
anchisiades. 


MATERIALS AND METHODS 


A cluster of 53 eggs was obtained by observing one female P. an- 
chisiades ovipositing on a 4-m high lemon (Citrus) tree at the edge of 
a grassy cattle pasture at “Finca La Tirimbina” at 13800 h on 4 March 
1982 in NE Costa Rica. This locality is about 10 km E of La Virgen 
(10°23'N, 84°07’W, 220 m elev.), Heredia Province, and well within 
the Premontane Tropical Wet Forest region (Holdridge 1967). The 
eggs were collected by cutting the branch with the leaf bearing them, 
and placing the cutting in a clean, air-tight, clear plastic bag. The 
larvae were reared following previously established methods (Young 
1972), which included daily observations and periodic changing of 
leaves and removal of frass and other debris. The duration of each life 
stage was measured, and feeding and resting behavior noted. The cul- 
ture of first instars was transported to Milwaukee, Wisconsin, where 
the rearing continued until adult emergence. During the Wisconsin 
rearing period, the larvae were fed leaves from a Citrus tree in the 
greenhouse at the Milwaukee Public Museum. 

The rearing period extended from 4 March through 23 April 1982, 
and during this time osmeterial secretions were collected from all avail- 
able larvae by instar. These secretions were collected in the standard 
way: each larva was gently pinched with fine forceps, and the everted 
osmeterium quickly wiped with a small square of filter paper and 
dropped immediately into a vial of methylene chloride. Several such 
“milkings’” were done within an instar between 1400-1500 h, and 
samples were thus obtained for the third through fifth instars. Milkings 
from different larvae were pooled at each sampling date as follows: 24 
third instars milked 1-2 April; 22 fourth instars milked 5-9 April; 22 
fifth instars milked 13-19 April. The apparency of odor associated with 
everted osmeteria was also noted. 


Chemical Analyses 


Gas chromatography-mass spectrometry subdivides complex com- 
pounds into molecular weight fractions. It was done using 15 m x 0.3 
mm I.D. OV-17 or SE-30 fused quartz capillary columns (J. and W. 
Scientific Co., Rancho Cordova, CA) in an LKB 2091 spectrometer, 
with a splitless injector system (J. and W. Scientific Co.). Confirmation 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of the low boiling esters was accomplished on an LKB 900 spectrometer 
using a 2 m X 2.5 mm I.D. 10% SP-1000 packed column. Synthetic 
octadecyl and eicosanyl esters were chromatographed on a 2 m X 2.5 
mm I.D. 1% OV-17 packed column. Both spectrometers were main- 
tained during scanning at 70 ev, at a source temperature of 270°, and 
at 270 amp ionizing current. 


Synthesis of Eicosanyl Esters of Valeric and Isovaleric Acids 


These compounds were prepared for use as standards by combining 
an excess (1 g) of the corresponding acid chlorides with 1 g of eicosanol 
in 8 ml of pyridine. After 1 h the mixtures were poured into water 
and extracted into ether, washed with dilute sodium bicarbonate, and 
the ether evaporated. The oils were then chromatographed directly 
providing only one peak on a 1% SE-54 packed column. 

Eicosyl valerate. Retention times, the result of component molecular 
weights and their percent representation in samples at a specified tem- 
perature, characterized specific complex compounds with mass spec- 
trometry. Retention temperature 270°, MS: m/z (rel. intensity) 382(0.9 
m*.), 353(0.4), 367(0.060), 340(0.4), 325(2), 280(8), 195(1), 181(1), 
167(2), 158(1), 153(2), 139(4), 125(7), 111(14), 103(100, valeric acid + 
H), 102(17), 97(24), 85(22), 83(23), 82(10), 71(13), 70(9), 69(14), 57(21), 
56(7), 43(6). 

Eicosyl isovalerate. Retention temperature 265°, MS: m/z (rel. in- 
tensity) 382(0.8, m*.), 367(0.4), 340(0.2), 325(1.5), 280(7), 252(8), 195(1), 
181(1), 167(2), 158(1), 153(8), 189(4), 125(6), 111(10), 103(100, valeric 
acid + H), 97(15), 85(19), 83(14), 71(7), 70(5), 69(9), 57(11), 48(4). 

Preparation of methyl! esters of osmeterial extract. Diazomethane 
in ether prepared from N-nitro-N-nitrosomethylguanidine (Aldrich 
Chemical Co., Milwaukee, WI) was added to 20 ul portions of the 
extract in methylene chloride until a yellow color persisted. Aliquots 
of this solution were directly injected. 


RESULTS 
Description of Early Stages 


Eggs (Fig. 1) spherical, sculptured, about 2 mm diam, with lateral 
pair of ridges fusing into bilobed knob; honey-colored; not changing 
in color before hatching; duration of stage: seven days. 

First instar cylindrical with fine down and slightly bulbous head; 
initially about 6 mm long; cuticle translucent amber, darkening to 
“dirty” greenish brown following first feeding on plant tissues; lateral 
body profile tapered; no tubercles and no discernible markings on cu- 
ticle; duration of stadium: seven days. 


VOLUME 40, NUMBER 1 





‘ g 
cs ee . wa > 
~~ . . : ; Q 
: ne ‘> ° A a 
: . ; WAS Bees 
ae 5 Vea Reo wy: % . 
: D Fass : hs 2S :- a 
‘ Pos a BS « be ‘ a 
sy Re ye Bs on er” ie a ee 


Fic. 1. Papilio anchisiades. (A) position of egg cluster on Citrus leaf (ventral surface); 
(B) orientation of individual eggs in cluster; (C) surface sculpturing of individual eggs; 


(D) second instars. 





Second instar (Fig. 1) similar to first but with larger head relative to 
trunk; more delineation of trunk segments; first three segments and 
last four dull orange, middle segments greenish; head glossy orange; 


attained body length of 10-13 mm in seven to nine days. 
Third instar (Fig. 2) strikingly different from previous instars; swelled 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Papilio anchisiades. (A) third instars immediately following molt; (B) fourth 
instars several days after molting; (C) fifth instar, lateral aspects; (D) aggregative behavior 
of fifth instars. 


thoracic region; coloration a variegated brown and white cuticle with 
“oily” appearance; head brownish orange and glossy; head hidden and 
overshadowed by anterior trunk region; attained a length of 20-23 mm 
in eight days. 


VOLUME 40, NUMBER 1 4] 


Fourth instar (Fig. 2) similar to third but with decrease in thoracic 
swelling; more pronounced mottling of rich brown and white blotches 
on trunk; both third and fourth instars with tubercles further described 
in Oliveira (1977); attained length of 32-36 mm in ten days. 

Fifth instar (Fig. 2) with trunk cuticle “lacework’ pattern of choc- 
olate-brown background with network of lines and blotches of white; 
cuticle “warty” due to small tubercles and foldings (further described 
in Oliveira 1977); prolegs white with brown speckling; head brownish 
and smaller than anterior trunk; dorsally trunk cuticle bears a series of 
diamondlike velvety-brown blotches; attained length of 59-62 mm in 
20 days. Total larval period: 45 days. 

All instars with deep-orange osmeteria, short and stubby in first three 
instars, long and filamentous in last two. Osmeterium of fifth instar 9- 
10 mm long. The prepupa (Fig. 3) contracted in body length and 
darkened in coloration before the final ecdysis. 

Pupa (Fig. 3) 37-39 mm long and 21-23 mm at greatest width; 
resembles broken twig; color pattern a variable mosaic of brown, gray, 
green, and white, but usually with large, “lichenlike’’ blotch on pos- 
terior two-thirds of wing pads extending posteriorly into dorsal area of 
abdomen; spiracle openings marked in black; duration of stage: 18-22 
days. Overall egg-to-adult time: 70-74 days. 

Adults eclosed rapidly, and wings were fully expanded (Fig. 3) with- 
in 25 min, and usually between 0800-0900 h. Sex ratio of 25 pupae: 
10 females and 15 males. 


Behavior of Stages 


Eggs placed in tight rows on the ventral surface of mature Citrus 
leaf (Fig. 1); even though “‘young” or “‘fresh” (greenish-yellow) leaves 
available, eggs were placed on older leaf, near the distal end of the 
branch; butterfly clung to edge of leaf and curled abdomen under 
while ovipositing for 1 h. When frightened away, it did not return to 
resume egg laying on the same or several subsequent days. Other ob- 
servations in Costa Rica indicate that this species oviposits on both 
mature (greenish-blue) leaves and yellowish-green fresh leaves of Cit- 
rus in both wet and dry forest regions. All eggs in the cluster touched 
one another and hatched synchronously taking about 4 h for all larvae 
to vacate egg shells. Egg shells were immediately devoured by larvae, 
and larvae remained as one group in the first two instars (Fig. 1). First 
instars occupied the same leaf as the eggs, and started feeding at the 
edge of the leaf (Fig. 4). Feeding throughout all instars was synchro- 
nous and nocturnal. Breakup into two or more subgroups began in the 
third instar and continued through the fifth (Fig. 2). Fourth instars 


42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 8. Papilio anchisiades. (A) prepupa; (B) pupa; (C) aggregation of pupae; (D) 
freshly eclosed adult. 


stayed in small groups on the branch rather than on leaves like previous 
instars. 

In field observations fourth and fifth instars aggregated on the trunk 
of the host, and pupation occurred on the trunk, on nearby buildings 
or other substrates near the host. 


VOLUME 40, NUMBER 1 43, 





Fic. 4. Papilio anchisiades. Feeding pattern of young larvae on a leaf of Citrus. 


Larvae of all five instars evert the osmeterium when prodded with 
forceps, but response is much quicker in the first three instars than in 
the last two. Eversion of the osmeteria in the first three instars was 
unaccompanied by odor at close range. A strong, disagreeable odor, 
best described as ‘‘sweaty socks,’ was apparent when the osmeteria of 
the last two instars were everted. Growth rates of larvae within a group 
were mostly similar, and molting was synchronous. Molting required 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


one to two days. However one to three individuals within the group 
were smaller, and differed by a full instar. Pupation appeared to occur 
in two “pulses,” with larger larvae (presumably females) being the last 
to pupate. 

In field observations, more than one cohort of larval P. anchisiades 
occurred in a single Citrus tree, often in trees with nests of paper 
wasps (Polistes and various polybiines). In a Citrus tree in a pasture 
at Palo Verde, Guanacaste Province, Costa Rica, studied between 9- 
11 November 1969, separate groups of 40-70 larvae were found, and 
each group contained 3-8 smaller larvae. Most larvae were fifth instar, 
and the smaller ones second and third instars. A total of nine pupae 
were scattered on branches, and all were being attacked by the ant 
Camponotus rectangularis Emery. An additional 25 pupae, without 
ants, were found on a nearby weather-beaten tool shed 5 m from the 
tree and separated from it by tall grasses. A total of 53 paper wasp 
nests were in the tree and another 74 nests on the shed. Individual 
wasps often perched near the pupae on the sides of the shed, but did 
not attack them. Pupae on the tree were not observed to be attacked 
by wasps. Successful eclosion of four adults was observed. No pupae 
on the shed were attacked by C. rectangularis during the two days of 
observation. 

Several third instars collected in October 1969 at Naranja, Zaragoza, 
El Salvador, were parasitized by the braconid Meteorus sp. A second 
unnamed species of the same genus has been recorded from P. an- 
chisiades in Venezuela (Paul Marsh, pers. comm.). Together, both rec- 
ords are new, and represent the first reports of parasitism by Meteorus 
on the Papilionidae (Paul Marsh, pers. comm.). 

A major feature of larval behavior in P. anchisiades in both field 
and laboratory is the close physical contact among individuals, al- 
though laboratory individuals sometimes rested and fed solitarily. 


Mass Spectral Analysis of Osmeterial Extracts 


Extracts of the fifth instar (Fig. 5) showed a poorly resolved series 
of short-chain acids and esters, followed by traces of sesquiterpenes 
eluting from 142-170°, and finally (Fig. 6) a series of hexadecyl, oc- 
tadecyl, and eicosanyl esters of butyric and valeric acids. The early 
eluting compounds were ethyl isobutyrate, methyl 2-methylbutyrate, 
ethyl 2-methylbutyrate, isobutyric acid, isovaleric acid and 2-methy]- 
butyric acid, eluting in that order. Reexamination on a 10% SP-1000 
packed column confirmed these assignments and revealed a trace of 
ethyl 3-hydroxybutyrate eluting just after ethyl 2-methylbutyrate. Also 
observed were traces of acetic acid and ethyl acetate. All compounds 
were identified by their mass spectra (Heller & Milne 1976). As found 


VOLUME 40, NUMBER 1 45 


Ethyl lsobutyrate | | | 
Isobutyric Acid 
Isovaleric Acid 
Ethyl 2-Methylbutyrate 


2-Methylbutyric Acid — 


RESPONSE 


ince Butyrate 
Octadecyl Valerate 


Eicosanyl Butyrate 


Eicosanyl Valerate 
Methyl 2-Methylbutyrate 
CoH ag 
Octadecanol CL | \ 
Va l 


249° 228° 205° 170° 142° 80° 50° 
TIME/TEMP 


Fic. 5. Chromatographic analysis of fifth-instar osmeterial secretion in Papilio an- 
chisiades (SE-30 capillary; 15 m x 0.30 mm I.D.; 10°/min.). 





by Honda (1981) for other Papilio species, 2-methylbutyric and iso- 
butyric acids were major components accompanied by smaller quan- 
tities of isovaleric acid. Conversion to the methyl esters allowed quan- 
titation of these acids in the ratio 1:0.75:0.021, respectively, as 
determined on an SE-54 capillary column. 

Expansion of the chromatographic region between 80° and 228° (Fig. 
6) allowed two terpenes, a-bergamotene and E-b-farnesene (peaks 1 
and 2), to be tentatively identified by comparison of their spectra with 
published compilations (Heller & Milne 1976). Peak 4 was tentatively 
identified as a-himachalene by similar comparison, while peak 9, as- 
sumed to be a sesquiterpene from its mass spectrum (Table 1), was 
unique to the fifth instar. The mass spectrum of peak 8 was similar 


46 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Mass spectra of sesquiterpenes identified in the osmeterial secretions of 
Papilio anchisiades. Retention temperatures (first numbers) and relative amounts ( ) refer 
to peaks in Fig. 7. 


1. a-bergamotene: m/z 204(M*., 2), 161(3), 131(5), 119(53), 93(87), 69(53), 55(40), 43(67), 
41(100) 

2. A teh isomer: m/z 204(M*., 5), 161(6), 183(14), 119(7), 93(41), 69(100), 55(20), 
43(23), 41(99) 

3. B-accoradiene: m/z 204(M*., 6), 182(2), 161(6), 183(16), 120(18), 107(8), 93(50), 91(25), 
81(25), 79(25), 69(100), 67(25), 57(20), 55(20), 41(90) 

4. “a-himachalene”’: m/z 204(M*., 25), 189(7), 161(10), 147(9), 188(17), 121(25), 119(22), 
93(100), 79(29), 69(21), 55(31), 48(63), 41(88) 

5. A farnesene isomer (same as 2 above, but mixed with a-himachalene) 

6. A farnesene isomer m/z 204(M*., 10), 161(18), 1383(20), 120(18), 109(16), 93(49), 
69(100), 41(79) 

7. “B-selinene”’: m/z 204(Mt., 15), 189(3), 161(3), 121(22), 119(20), 109(21), 98(100), 
80(80), 69(25), 41(42) 

8. Unknown: m/z 220(M*., 20), 205(5), 177(4), 163(2), 151(6), 149(4), 137(100), 185(53), 
110(76), 109(51), 95(46), 82(29), 69(35), 55(40), 43(20), 41(85) 

9. Unknown (only in 5th instar): m/z 204(M*., 25), 189(18), 169(31), 183(25), 121(50), 
119(44), 105(52), 93(69), 91(50), 79(88), 77(31), 69(25), 55(44), 53(38), 43(81), 41(100) 


but not identical to the spectrum of caryophyllene oxide reported by 
Honda (1981). No evidence was found for monoterpenes or the ele- 
mene, selinene or germacrenes reported by Honda (1981). Also iden- 
tified in this sean were C,,-C,, saturated and unsaturated hydrocarbons 
as well as naphthalene, dichlorobenzene, and phthalates, all of which 
are regarded as artifacts. | 

The acid components of the hexadecyl, octadecyl, and eicosy] esters 
were expected to be isobutyric and either 2-methylbutyric or isovaleric 
acids in view of the large quantities of the corresponding free acids 
that were present (Fig. 5). In fact, comparison of the mass spectra of 
the last peak with spectra of synthesized samples of eicosyl n-valerate 
and eicosyl isovalerate reveals that the natural product is the former 
ester. Thus, the molecular ion of eicosyl isovalerate was slightly less 
intense relative to high mass peaks, and showed enhanced loss of meth- 
yl compared to eicosyl valerate. The other three peaks are also esters 
of the n-butyric and n-valeric acids by the same reasoning. Fig. 6 also 
shows the presence of the corresponding alcohols, octadecanol and ei- 
cosanol, easily identified by reference to library spectra. 

Gas chromatograms of the third and fourth instars were nearly iden- 
tical, but presented an entirely different picture (Fig. 7). Both short 
and long chain acids and esters were missing, and only sesquiterpenes 
were present. As in extracts from the fifth instar, only a-bergamotene, 
-acoradiene and three farnesene isomers were identified with confi- 
dence by comparison with reference spectra. The major component 
was a compound whose spectrum resembled, but was not identical 


VOLUME 40, NUMBER 1] 47 








Octadecanol 
8 
C,H,  Dibutylphthalate 
LW 
7) 
Zz 
oO 
Diisobutylphthalate = 
n-C,,H,,  Diethylphthalate cc 
Eicosanol C_H 
ly WS ain 
n-C..H n-C,.H3, 
cae | Oaeo: : 
n-C.H ~ | ZA i 
22° 46 aS | AAC ee 
Za) 


IL 


Hexadecy] butyrate 





Hexadecyl valerate | 





249° 228° 190° 170° 150° 142° 80° 5Oe 


TIME/TEMP 


Fic. 6. Mass spectral analysis of fifth-instar osmeterial secretion in Papilio anchisi- 
ades, highlighting the terpene region of the spectra (SE-30 capillary; 15 m x 0.3 mm 
I.D.; 10°/min.). 


with, caryophyllene oxide as reported by Honda (1981). Peaks 4 and 
7 are very similar, and both resemble library spectra (Heller & Milne 
1976) of a-himachalene or b-selinene, but neither corresponds to the 
b-selinene spectrum reported by Honda (1981). Mass spectra of ses- 
quiterpenes are shown in Table 1. 


DISCUSSION 


Papilio anchisiades, along with P. cresphontes and P. thoas, and a 
few others, exploits various Rutaceae as larval food plants (Brower 
1958). It differs from other rutaceous-feeding Papilio species by its 
unique larval aggregative habits, a result of cluster egg placement on 
the larval host plants. The rutaceous-feeding habit is shared world- 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


wide by several species of Papilio (Munroe 1960), an association per- 
haps mediated by the rich profiles of specific alkaloids so characteristic 
of this plant group (Hegnauer 1963). We propose that the widespread 
abundance of P. anchisiades from southern Brazil to Mexico and even 
southern Texas (Ehrlich & Ehrlich 1961), is in part due to the broad 
distribution of Citrus as an exotic rutaceous host plant coupled with 
the distribution of other Rutaceae with forest habitats. 

Larval host-plant selection in butterflies usually involves a highly 
structured sequence of visual and olfactory responses (Crane 1955, 
Swihart & Swihart 1970, Ilse 1937). Vaidya (1969) studied the color 
preferences of ovipositing P. demoleus L. on Citrus in laboratory stud- 
ies, and concluded that both color and scent are required for proper 
egg placement and noted that butterflies preferred blue-green hues of 
leaves over yellow-green hues. Within the Rutaceae, differences among 
genera and species for certain substances in the leaves determine pat- 
terns of host specificity in egg-placement behavior among different 
papilionids (Ichinose et al. 1981). Age-specific differences in odor and 
color in Citrus and other Rutaceae may determine patterns of egg 
placement in P. anchisiades in nature, and the field data for this species 
in Costa Rica supports a partial preference concept for the older, more 
bluish-green leaves of host plants. Depending on annual phenological 
patterns of flushing, such egg-placement substrates may vary in abun- 
dance at a locality and influence the abundance of the butterfly pop- 
ulation, or result in oviposition on more yellowish green leaves. In a 
highly seasonal lowland area such as Guanacaste, such effects might 
be even more pronounced than in less seasonal Atlantic zone habitats 
in Costa Rica and elsewhere in southern Central America. Ross (1964b) 
noted that P. thoas autocles Rothschild & Jordan frequently oviposits 
on fresh leaves of the larval host plants, including Piper spp. (Pipera- 
ceae) and Citrus in Mexico. Papilio aristodemus ponceanus oviposits 
on young shoots of Zanthoxylum fagara (Rutaceae), “Wild Lime,” and 
first instars readily devour the young leaves without difficulty (Rut- 
kowski 1971). Tough, thick leaves of Rutaceae used by Papilio species 
may retard normal growth and development of caterpillars (Watanabe 
1982), thereby selecting for egg placement on young, tender leaves. 

Several studies reveal that the attraction of parasitoids to their phy- 
tophagous hosts is often mediated by the aromatic substances emitted 
by the host plant (Herrebout 1969, Read et al. 1970). We suspect that 
Papilio species associated with the highly aromatic Rutaceae are sub- 
ject to such parasitism, and several larvae within an aggregate of P. 
anchisiades can be killed by the braconid Meteorus sp. Such interac- 
tions may extend to predatory arthropods such as the ant C. rectan- 
gularis associated with Citrus in lowland Guanacaste, even though 


VOLUME 40, NUMBER 1 49 


predation by paper wasps under the same conditions may be minimal 
or nonexistent. 

The size of larval groups of P. anchisiades in Citrus varies, and the 
group observed in the present study might have been small since the 
ovipositing butterfly was frightened away. The larvae have been noted 
to defoliate a tree (Caracciolo 1891), and very large groups of larvae 
have been found on individual trees (Moss 1919). Pupae are often 
found on various substrates away from the host tree (Moss 1919), and 
the present study suggests that mortality from at least one ant species 
might be less for pupae off the host tree than for those remaining on 
1 

Although the disagreeable odor from the osmeteria of the older lar- 
vae is well known (Stoll 1781, Carracciolo 1891, Moss 1919), the func- 
tional role of the secretion remains unknown, although the components 
are defensive against ants (Honda 1983). The precise egg-placement 
behavior of P. anchisiades suggests that the species is a specialist on 
Rutaceae, a condition that further suggests coevolved associations with 
parasitoids and predators that cue into the aromatic properties of Cit- 
rus and other genera within the family. The cryptic appearance and 
behavior of larvae of all instars, and the cryptic appearance of the 
pupa, suggest that this species is palatable to visually foraging predators 
such as lizards and birds (Brower & Brower 1964). When this first-line 
defense is penetrated by an attacker, the odor defense associated with 
the osmeterium might be used to thwart attack (Eisner & Meinwald 
1965, Honda 1983). All rutaceous-feeding Papilio species appear to 
have cryptic coloration and habits (Munroe 1960). 

We suggest that aggregative behavior in the larval stages of P. an- 
chisiades enhances visual crypsis to some predators such as birds and 
lizards. The combined aggregate of several fifth instars on the bark of 
the host tree creates the image of a mottled blotch of false lichens and 
bark on the trunk. Similarly, the tightly packed clusters of younger 
larvae on the ventral surfaces of Citrus leaves resemble dead or dying 
plant tissue destroyed by a pathogenic microorganism. A large aggre- 
gation of fifth instars positioned at the junction between the trunk and 
main branches of a Citrus tree in Trinidad resembled a “‘clot of wet 
feces” to both L. P. Brower and P. M. Shepard (L. P. Brower, pers. 
comm.). Aggregative behavior of the larvae, however, may not deter 
predation by birds. On 28 July 1986 one of us (A.M.Y.) observed an 
unidentified jay-size bird pluck off a Citrus leaf bearing 50 young 
third-instar P. anchisiades (at 0530 h) at “Finca La Lola” in Costa 
Rica. The bird then devoured all the larvae in a few seconds. 

The osmeterial secretions from third and fourth instars of P. an- 
chisiades are similar to those of other Papilio species in being domi- 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


® 


y, B-Farnesene 
Isomer 


® 


"‘a-Himachalene”’ 
Raed 


RESPONSE 


Farnesene 


Isomers \ 


©) 
@ a) CL 


“B-Selinene”’ 
ra “‘a-Bergamotene”’ 


250° 170° = 140° 90° 


a-Acoradiene 


TIME / TEMP 


Fic. 7. Mass spectral analysis of fourth-instar osmeterial secretion in Papilio anchisi- 
ades, highlighting sesquiterpene region of the spectra. 


nated by terpenes. Burger et al. (1978) and Honda (1980a, b, 1981) 
reported that the secretions of earlier instars of several Papilio species 
are made up of mono- and sesquiterpenes. While we did not detect 
monoterpenes in the secretions of P. anchisiades, at least seven ses- 
quiterpenes fortify the osmeterial exudate (Fig. 7). Honda (1981) pre- 
viously identified sesquiterpenes in the osmeterial secretions of five 
species of Papilio, but several of those produced by P. anchisiades 
appear to be different from those produced by the Japanese species. 
Earlier investigations established that the osmeterial secretions of a 
variety of Papilio, Baronia, and Eurytides species were dominated by 
isobutyric and 2-methylbutyric acids (Eisner & Meinwald 1965, Cross- 
ley & Waterhouse 1969, Eisner et al. 1970, Burger et al. 1978, Lopez 
& Quesnel 1970). However, it was subsequently demonstrated that this 


VOLUME 40, NUMBER 1 51 


acidic duet is characteristic of the osmeterial secretions of the fifth 
instar. In contrast, the secretions of earlier instars of several Papilio 
species lack the short-chain acids produced by fifth instars, and a va- 
riety of terpenes are produced by younger larvae (Burger et al. 1978, 
Honda 1980a, b, 1981). 

The osmeterial secretion of the fifth instar of P. anchisiades contains 
isobutyric and 2-methylbutyric acids, but, in addition, isovaleric acid, 
a compound detected as a minor osmeterial constituent in two other 
Papilio species (Honda 1981). Although isobutyric and 2-methylbutyr- 
ic acids have been encountered as the acidic moieties of short-chain 
esters in the osmeterial secretions of P. anchisiades and other species 
(Burger 1978, Honda 1981), long-chain esters containing butyric and 
valeric acids (Fig. 5) have not been reported previously from papilionid 
osmeterial secretions. Thus, fifth-instar larvae of P. anchisiades are 
distinctive in producing osmeterial secretions containing esters such as 
hexadecyl! valerate (Fig. 6) and octadecyl butyrate (Fig. 5). It is not 
clear why the dominant free acids in the secretion—isobutyric and 
2-methylbutyric—have not been utilized as the acid moieties of these 
long-chain esters. 

Sesquiterpenes in the osmeterial secretion of the fifth instar is un- 
usual, since this class of compounds has been identified in the secretions 
of earlier instar Papilio (Burger et al. 1978, Honda 1980a, b, 1981). 
However, one sesquiterpene has been identified in the secretion of P. 
protenor (Honda 1980), and three in that of P. memnon (Honda 1981). 
Papilio anchisiades is unusual in having almost as many sesquiterpenes 
(five) in the secretion of the fifth instar as in that of earlier instars 
(seven). 

The secretions of the last instar of P. anchisiades differs from those 
of any Papilio species similarly analyzed in containing aliphatic hy- 
drocarbons and long-chain alcohols (Fig. 7). Nine aliphatic hydrocar- 
bons are present, and these are accompanied by C,, and C,, alcohols. 
With the presence of esters such as hexadecy] valerate, the distinctive- 
ness of this osmeterial secretion is further evident. 

Previous investigators demonstrated that the secretions of younger 
larvae were qualitatively richer than those of fifth instars (Burger et 
al. 1978, Honda 1980a, b, 1981); the opposite is true for the secretions 
of earlier and fifth instars of P. anchisiades. Whereas the third and 
fourth instar secretions contain seven sesquiterpenes (Fig. 7), the fifth 
instar secretion contains more than 30 compounds (Figs. 5, 6) including 
acids, hydrocarbons, esters, alcohols, and sesquiterpenes. Qualitatively, 
the fifth-instar secretion of P. anchisiades exceeds that known for any 
instars of any Papilio species. 

If it is assumed that the chemical (osmeterial) defenses of earlier- 
instar Papilio evolved as deterrents against predators different than 


52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


encountered by the fifth instar, then the differences in the chemistry 
of these instars is explicable. There is little specific evidence on what 
organisms constitute enemies for larvae of P. anchisiades of any instar. 
Whatever the selection pressures were for producing the fifth-instar 
exudate, they have resulted in the most diverse osmeterial secretion 
encountered in the genus Papilio. 

The greater heterogeneity and complexity of the osmeterial secretion 
of fifth instars in P. anchisiades suggests that the system becomes most 
functional in this instar. The pungent odor emitted in the fifth instar 
results from isobutyric and 2-methylbutyric acids, which are lacking 
in the earlier instars. The occurrence of some components (sesquiter- 
penes) of earlier-instar osmeterial secretions in the fifth instar indicates 
that the biochemical pathways underlying the synthesis of these sub- 
stances are not completely turned off in the fifth instar. Both qualitative 
and quantitative changes figure in the regulation of secretion in the 
fifth instar of P. anchisiades. 

Our results are largely due to the application of capillary-column 
gas chromatography, which enabled detecting of minute amounts of 
specific components in the fifth instar, substances that might have been 
overlooked otherwise. 


ACKNOWLEDGMENTS 


Field work in Costa Rica was supported by N.S.F. Grant GB-7805 (D. H. Janzen, 
principal investigator) in 1969-70; N.S.F. Grant GB 33060, Friends of the Milwaukee 
Public Museum, and the American Cocoa Research Institute. We thank Thomas Emmel 
and Michael Salmon for fruitful discussions, and E. O. Wilson for identifying the ant 
species. Susan Borkin assisted in rearing and milking the larvae. Special thanks to A. 
Muyshondt for providing the specimens and record of larval parasitism from El Salvador. 
Paul Marsh identified the braconid wasp and shared unpublished data with us. We thank 
L. P. Brower for helpful suggestions on the manuscript. 


LITERATURE CITED 


BROWER, L. P. 1958. Larval foodplant specificity in butterflies of the Papilio glaucus 
group. Lepid. News 12:103-114. 

BROWER, L. P. & J. V. Z. BROWER. 1964. Birds, butterflies, and plant poisons: A study 
in ecological chemistry. Zoologica 49:137-159. 

BurRGER, B. V., M. ROTH, M. LERoux, H. S. C. Spies, V. TRUTER & H. GEERTSMA. 
1978. The chemical nature of the defensive larval secretion of the citrus swallowtail, 
Papilio demodocus. J. Insect Physiol. 24:803-810. 

CARRACCIOLO, H. 1891. Notes and news. Entom. News 2:52. 

CRANE, J. 1955. Imaginal behavior of a Trinidad butterfly, Heliconius erato hydara 
Hewitson, with special reference to the social use of color. Zoologica 40:167-196. 

CrossLEy, A. C. & D. F. WATERHOUSE. 1969. The ultrastructure of the osmeterium 
and the nature of its secretion in Papilio larvae (Lepidoptera). Tissue and Cell 1: 
525:554. 

DewiTz, H. 1878. Entwickelung einiger Venezuelanischer Schmetterlinge nach Beo- 
bachtungen von Gollmer. Arch. Naturgesch. 44:1-36. 

EHRLICH, P. R. & A. H. EHRLICH. 1961. How to know the butterflies. Brown, Dubuque, 
Iowa. 262 pp. 

EISNER, T. & Y. C. MEINWALD. 1965. Defensive secretion of a caterpillar (Papilio). 
Science 150:1733-1735. 


VOLUME 40, NUMBER 1 53 


EISNER, T., T. E. PLISKE, M. IKEDA, D. F. OWEN, L. VASQUEX, H. PEREZ, J. G. 
FRANCLEMONT & J. MEINWALD. 1970. Defensive mechanisms of arthropods— 
XXVII. Osmeterial secretion of papilionid caterpillars. Ann. Entom. Soc. Am. 63: 
914-915. 

HEGNAUER, R. 1963. The taxonomic significance of alkaloids, pp. 289-427. In Swain, 
T. (ed.), Chemical plant taxonomy. Academic Press, London & New York. 

HELLER, S. & G. W. A. MILNE, eds. 1976. EPA/NIH mass spectral data base. Publ. 
NBS 20402 NSRDS-NB563, U.S. Govt. Printing Office, Washington, D.C. 

HERREBOUT, W. M. 1969. Some aspects of host selection in Eurarcela rutilla Vill. 
(Diptera: Tachinidae). Netherlands J. Zool. 19:1-104. 

Honpa, K. 1980a. Volatile constituents of larval osmeterial secretions in Papilio pro- 
tenor demetrius. J. Insect Physiol. 26:39—45. 

1980b. Osmeterial secretions of papilionid larvae in the genera Luehdorfa, 

Graphium and Atrophaneura. Insect Biochem. 10:583-588. 

1981. Larval osmeterial secretions of the swallowtails. J. Chem. Ecol. 7:1089- 

11138. 

1983. Defensive potential of components of the larval osmeterial secretion of 
papilionid butterflies against ants. Physiol. Entom. 8:173-179. 

ICHINOSE, T., H. ABE & Y. TERAMOTO. 1981. Relationship between host plant ranges 
of three papilionid butterflies and oviposition-inducing contact chemicals in their 
host plants. Appl. Entom. Zool. 16:493-496. 

ILsE, D. 1937. New observations on responses to colours in egg-laying butterflies. Nature 
140:544. 

JONES, C. 1882. Metamorphoses of Brazilian Lepidoptera. Proc. Lit. Philos. Soc., Liv- 
erpool 36:327-377. 

Lopez, A. & V. C. QUESNEL. 1970. Defensive secretions of some papilionid caterpillars. 
Carib. J. Sci. 10:5-7. 

Moss, A. M. 1919. The Papilios of Para. Novitat. Zool. 26:259-319. 

MUNROE, E. G. 1960. The generic classification of the Papilionidae. Can. Entom., 
Suppl. 17:1-51. 

OLIVEIRA, B. L. DE. 1977. Contribuicaeo ao conhecimento da biologia de Papilio 
anchisiades capys (Hubner, 1809) (Lepidoptera-Papilionidae). Dusenia 10:89-95. 

READ, D. P., P. P. FEENy & R. B. Root. 1970. Habitat selection by the aphid parasite 
Diaeretiella rapae (Hymenoptera: Braconidae) and hyperparasite Charips brassicae 
(Hymenoptera: Cynipidae). Can. Entom. 102:1567-1578. 

Ross, G. N. 1964a. Life history studies on Mexican butterflies. I. Notes on the early 
stages of four papilionids from Catemaco, Veracruz. J. Res. Lepid. 3:9-18. 

1964b. Life history studies on Mexican butterflies. II]. Nine Rhopalocera (Pa- 
pilionidae, Nymphalidae, Lycaenidae) from Ocotal, Chico, Veracruz. J. Res. Lepid. 
3:207-229. 

RUTKOWSKI, F. 1971. Observations on Papilio aristodemus ponceanus (Papilionidae). 
J. Lepid. Soc. 25:126-136. 

SEITZ, A. 1908. Macrolepidoptera of the World. Vol. 5. American Rhopalocera. A. 
Kernan, Stuttgart. 

STOLL, C. 1781. In Pieter Cramer (ed.), De uitlandsche Kapellen voorkomende in de 
drie Waereld-Deelen Asia, Africa, en America. Suppl. 3, Amsterdam. 384 pp. 
SELIGMAN, I. M. & F. A. Doy. 1972. -Hydroxy-n-butyric acid in the defensive secretion 

of Papilio aegus. Comp. Biochem. Physiol. 41:341-342. 

SWIHART, C. A. & S. L. SwIHART. 1970. Colour selection and learned feeding prefer- 
ences in the butterfly, Heliconius charitonius Linn. Anim. Behav. 18:60-64. 

VaIpyA, V. G. 1969. Investigations on the role of visual stimuli in the egg-laying and 
resting behaviour of Papilio demoleus L. (Papilionidae, Lepidoptera). Anim. Behav. 
17:350-355. 

WATANABE, M. 1982. Leaf structure of Zanthoxylum ailanthoides Sieb. et Zucc. (Ru- 
tales: Rutaceae) affecting the mortality of swallowtail butterfly, Papilio xuthus L. 
(Lepidoptera: Papilionidae). Appl. Entom. Zool. 17:151-159. 

YounNG, A. M. 1972. Breeding success and survivorship in some tropical butterflies. 
Oikos 23:318-326. 














Journal of the Lepidopterists’ Society 
40(1), 1986, 54 


GENERAL NOTE 


SATURNIA WALTERORUM (SATURNIIDAE) IN MEXICO: 
A NEW NATIONAL RECORD 


Until now Saturnia walterorum (Hogue & Johnson) had not been taken in Mexico. It 
had been known to occur only in San Diego, Los Angeles, and Orange counties of 
California. A previous reference to specimens captured in San Luis Obispo Co. (Tilden 
1945, Pan-Pac. Entomol. 21:32-33) is in error, as the cited specimens were examined by 
Tuskes and Collins (1981, J. Lepid. Soc. 35:1-21) and found not to be typical walterorum. 

Both male and female are diurnal, and are not attracted to light. The seasonal flight 
period appears restricted to a few of the warmest days between late February and early 
June (Tuskes 1974, J. Lepid. Soc. 28:172-174). The insect is not abundant, and is easily 
overlooked. 

Our experience with this species in coastal San Diego Co. suggests the peak daily flight 
period for males is mid-morning, diminishing greatly before noon. 

On 1 April 1985 at 1130 h, we placed two newly emerged captive-reared females of 
this species in a screen cage among chaparral near Ensenada, Baja California, Mexico, 
about 100 km south of the United States border. The small amount of natural vegetation 
at this site was similar to that of areas near San Diego, California, and included Rhus 
laurina (Nuttall) and species of Ceanothus, Rhamnus and Adenostoma. Rhus laurina 
appears to us to be the preferred food plant of Saturnia walterorum in the coastal areas 
of its range. ; 

At 1140 h a single male was attracted to the calling females and was captured. No 
additional males had appeared by 1300 h, at which time we left the area. 

The captured specimen was placed in the San Diego Natural History Museum, San 
Diego, California. 


KirnBY L. WOLFE AND MARVIN D. VALVERDE, Route 5, Box 169-C, Escondido, Cali- 
fornia 92025. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 55-58 


PYRGUS COMMUNIS AND P. ALBESCENS (HESPERIIDAE) 
IN NEVADA 


GEORGE T. AUSTIN 


Nevada State Museum and Historical Society, 
700 Twin Lakes Drive, Las Vegas, Nevada 89107 


ABSTRACT. Based on more than 500 male genitalia, the Pyrgus communis phe- 
notype replaces the P. albescens phenotype latitudinally and elevationally in Nevada. 
Intermediates are known where their distributions meet and overiap. 


The status of Pyrgus communis (Grote) and Pyrgus albescens Plotz 
(Hesperiidae: Pyrginae) has been in question up to the present. They 
have been treated as separate species, as subspecies, or neither (Tilden 
1965). Even the most recent regional and taxonomic treatments vary. 
They were considered subspecies of P. communis by Stanford (in Fer- 
ris & Brown 1981) but as full species by Miller and Brown (1981). The 
two taxa are often segregated by ecology and geography but there are 
areas of sympatry or near sympatry in southwestern United States and 
adjacent Mexico. In some latter areas, intermediates are known (Tilden 
1965). In others, they are said to occur in close proximity, but no 
mention is made of intermediates (Ferris 1976, Stanford in Ferris & 
Brown 1981, Holland 1984); some workers have never seen an inter- 
mediate (Ferris, H. A. Freeman, pers. comm.). The present paper sum- 
marizes their status and distribution in Nevada. 

More than 500 male adults from Nevada in the Nevada State Mu- 
seum and in the author’s collection were examined. The left valva of 
each was classified into one of three configurations, the variations of 
which are indicated in Fig. 1. These were assigned to P. albescens, P. 
communis, and intermediate, and their distributions were mapped. 

The valvae of individuals assigned to nominate P. communis have 
a long and recurved dorsal process terminating in two sharply pointed 
prongs (Fig. 1). The lengths of the dorsal process and the prongs vary. 
On some individuals, one of the prongs is shorter than the other; on 
most they are equal. The valvae of individuals assigned to P. albescens 
have no dorsal process but usually have a single, short prong anterior 
to the tip (Fig. 1). Intermediates show various degrees of development 
in the dorsal process and the double prongs (Fig. 1). There was no 
difference in wing pattern between the genitalic phenotypes; their 
seasonal variation is likewise identical. 

Individuals of the P. communis phenotype occur throughout Ne- 
vada (Fig. 2); those of the P. albescens and intermediate phenotypes 
occur in southern Nevada except for one P. albescens from Carson 
City (Fig. 2). At most stations where P. albescens were taken, inter- 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


> oe 


communis 


5D De 


intermediates 


a/bescens 


Fic. 1. Variation in the left valvae of Pyrgus communis in Nevada. 


mediates and P. communis were taken also. Individuals with inter- 
mediate valvae occur only within the range of P. albescens. There is 
no strict ecological or elevational segregation in southern Nevada, but 
phenotype proportions do vary. The P. albescens phenotype dominates 
at lower elevations and latitudes. Intermediates and P. communis be- 
come more prominent with increase in elevation and latitude (Table 
1, Fig. 2). In the Newberry Mountains, Las Vegas Valley, and the lower 
slopes of the Spring Mountains, P. albescens accounts for more than 
60% of the individuals, and P. communis for less than 6%. At moderate 
elevations of the Spring Mountains, there is an increase in the P. com- 
munis phenotype and at the higher elevations and in Moapa Valley, 
intermediates predominate. 

The Nevada distribution is compatible with that previously noted 


TABLE 1. Proportion of P. albescens, P. communis and intermediate phenotypes 
from different locations in southern Nevada. 


P Inter- P 


Location albescens mediate communis N 
Newberry Mountains (<1,200 m) 60 36 4 25 
Las Vegas Valley (600-900 m) 62 33 5 2) 
Low slopes, Spring Mts. (<1,500 m) 65 29 6 17 
Mid elevations, Spring Mts. (1,500- 
2,100 m) 57 24 19 84 
High elevations, Spring Mts. (>2,100 m) 20 60 20 15 


Moapa Valley 34 48 18 91 


VOLUME 40, NUMBER 1 57 





e@ communis 
o a/bescens 

w intermediates 
A both 


A both and intermediates 
WV communis and intermediates 
V a/bescens and intermediates 


Fic. 2. Distribution of Pyrgus communis in Nevada. 


(Tilden 1965) for Pyrgus communis; the latter is a more northern and 
higher elevation phenotype, P. albescens, a lower-elevation and more 
southerly phenotype. Intermediacy, at least in southern Nevada, is 
greater than previously reported. This indicates that the two pheno- 
types are closely related, and are probably no more than allopatric 
subspecies of Pyrgus communis. 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


I thank H. A. Freeman and C. D. Ferris for their comments on these taxa. Thanks are 
also due two anonymous reviewers for their useful comments on the manuscript. 


LITERATURE CITED 


FERRIS, C. D. 1976. A checklist of the butterflies of Grant County, New Mexico and 
vicinity. J. Lepid. Soc. 30:38—49. 

FERRIS, C. D. & F. M. BRown. 1981. Butterflies of the Rocky Mountain States. Univ. 
Oklahoma Press, Norman. 

HOLLAND, R. 1984. Butterflies of two northwest New Mexico mountains. J. Lepid. Soc. 
38:220-234. 

MILLER, L. D. & F. M. BRown. 1981. A catalogue/checklist of the butterflies of 
America north of Mexico. Lepid. Soc. Mem., No. 2. 

TILDEN, J. W. 1965. A note on Pyrgus communis and Pyrgus albescens (Hesperiidae). 
J. Lepid. Soc. 19:91-94. 


ANNOUNCEMENT 


INAUGURATION OF MANUSCRIPT DATING IN THE JOURNAL 


Received and accepted dates will appear at the end of all research reports published 
in the Journal starting with submissions received in 1986. Such dating is practiced by 
many scholarly journals. It has at least three purposes. First, it encourages editors, re- 
viewers, and authors to speed manuscript processing. Second, it tells prospective contrib- 
utors how long manuscript processing might take. Third, it enables more accurate dating 
of ideas should issues of history or priority arise. 

To better serve these purposes, new received dates may be assigned to some revised 
manuscripts. Examples are those received more than two years after the editor requests 
them, and those with excessively broadened scopes. 

Received and accepted dates should make the Journal more useful to readers and 
authors alike. 


WILLIAM E. MILLER, Editor 


Journal of the Lepidopterists’ Society 
40(1), 1986, 59-63 


FIRST REPORTED MALES, SPECIES STATUS, AND 
AFFINITIES OF EPARGYREUS SPANNA 
EVANS (HESPERIIDAE) 


KURT JOHNSON 


Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024 


DAVID MATUSIK 


Department of Entomology, Field Museum of Natural History, 
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605 


ABSTRACT. Two males of E. spanna, an Hispaniolan endemic formerly reported 
from only two females, were collected in Pedernales Province, Dominican Republic, in 
1985. Male genitalia and other characters support species status of E. spanna but not its 
long-supposed affinity to E. antaeus Hewitson, endemic to Jamaica, and to E. zestos 
Geyer of trans-Caribbean distribution. Genitalically, E. spanna resembles E. windi Free- 
man of Mexico, E. aspina Evans of Colombia and E. tmolis Burmeister of western 
Argentina. Apparent affinities of E. spanna and some other recently described Antillean 
endemics do not support the island subspecies view of Caribbean biogeography. These 
taxa evidence most immediate kinship to particular mainland taxa, not most geograph- 
ically proximate Antillean congeners. 


Epargyreus spanna Evans (1952) has hitherto been known only from 
the holotype female (labelled “Santo Domingo” [Dominican Republic 
(DR)]) in the British Museum (Natural History), and a second female 
reported by Gali and Schwartz (1983) from west of Jayaco, La Vega 
Province, DR in the Albert Schwartz collection (Miami, Florida). 

Brown and Heineman (1972) suggested that E. spanna might rep- 
resent a subspecies of E. antaeus Hewitson. The latter is endemic to 
Jamaica, and among Antillean Epargyreus, shares with E. spanna the 
bright silver-white undersurface stripe on the hindwing. Riley (1975) 
figured both species, and Gali and Schwartz (1983), on the basis of 
wing character comparison, considered the two different species, pend- 
ing examination of E. spanna males. Both Evans and Freeman (1969, 
1977) emphasized the importance of male genitalic characters in dif- 
ferentiating Epargyreus taxa. 

In 1985, we collected extensively in the DR and, in Pedernales Prov- 
ince, collected two female and two male E. spanna. The precise lo- 
calities of these collections are: (1) a female of 830 mm forewing length 
(base to apex) taken between 0930 and 1200 h (EDT) on 30 June 1985 
at 1,350 m altitude, Aceitillar, 12 km NW of Las Abejas, Pedernales 
Province, DR, on a mountain path, in mesic broad-leaf deciduous for- 
est, in sunny weather (in David Matusik Collection [DMC)); (2) a fe- 
male (not measured) taken at same location at about the same time on 
1 July 1985 (in the Museo Nacional de Historia Natural, Santo Domin- 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. “Designated allotype” male of Epargyreus spanna. Left: upper surface; right: 
undersurface. 


go); (3) a male of 28 mm forewing length taken between 0930 and 
1200 h (EDT) on 4 July 1985 at 1,250 m altitude, about 5 km NNW 
of the locality cited in (1) in mesic broad-leaf deciduous forest, in partly 
sunny weather (DMC); and (4) a male of 29 mm forewing length taken 
at the same location at about the same time on 8 July 1985. Both 
females were collected while flying about 13 cm above the ground in 
zigzag flight, which alternated with periods of alighting on patches of 
bare ground usually about 4 m apart. Both males were collected while 
perched on broad-leaf deciduous foliage about 2.4 m above the ground, 
between leaf-to-leaf flights. The flight of males between leaves was 
notably slower than that of the females between patches of bare ground. 
The dates cited above, along with that of Gali and Schwartz [17 Au- 
gust], indicate the flight period of E. spanna to be at least three weeks. 

Considering the apparent rarity of E. spanna, and for reference 
purposes, we follow Smith (1983) and designate the last-cited male as 
“designated allotype”. The specimen, marked as such, is in the collec- 
tion of the American Museum of Natural History (AMNH), and is 
illustrated here (Fig. 1). The category “designated allotype’” has no 
status according to the International Commission on Zoological No- 
menclature Code, but is viewed as having diagnostic utility (Frizzell 
1933, Gloyd 1982, Smith 1983). 

Collection of the E. spanna male enabled examination of its genitalia 
(Fig. 2). Comparison of the genitalia with other Epargyreus taxa avail- 
able to us in AMNH genitalic material of W. H. Evans and H. A. 


VOLUME 40, NUMBER 1 61 


. 

> 

2 

S 

Py 

‘6 

i 

& 

cae at 
yf 
few 





Fic. 2. Male genitalia of Epargyreus spanna and E. antaeus. Above right: Dorsal view 
of E. spanna uncus and tegumen. Above left: Lateral view of E. spanna genitalia with 
aedeagus removed. A, Broad cephalad area; B, Reduced, cephalad-located dorsal process; 
C, Broadly thickened terminus with wide-based terminal hook. Immediately beneath: 
Lateral view of E. spanna aedeagus. Bottom: E. antaeus. D, Lateral view of valve; E, 
Lateral view of uncus; F, Lateral view of saccus; G, Lateral view of aedeagus; H, Dorsal 
view of tegumen and uncus. 


Freeman indicates that E. spanna is specifically distinct from its con- 
geners. These taxa are spina Evans, aspina Evans, antaeus, orizaba 
Scudder, exadeus Hiibner, windi, cruza Evans, deleoni Freeman, spi- 
nosa Evans, clarus Cramer, brodkorbi Freeman, zestos, tenda Evans, 
plus additional taxa studied from figures in Evans (1952). The genitalia 
do not reflect the close relation claimed for E. spanna, E. antaeus and 
E. zestos by Brown and Heineman (1972) and Riley (1975). Although 
E. spanna is similar to E. antaeus on the wing undersurface, the gen- 
italia of E. spanna are most like E. windi Freeman (type locality [TL] 
Ajijic [Jalisco], Mexico) and also similar to E. aspina Evans (TL Bogota, 
Colombia) and E. tmolis Burmeister (TL Buenos Aires, Argentina). 
Along with E. spanna, the above three taxa have on the male valvae 
a broad cephalad area (Fig. 2, A), a reduced and cephalad located 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


dorsal process (Fig. 2, B), and a broadly thickened terminus with wide- 
based terminal hook (Fig. 2, C). Epargyreus zestos and E. antaeus 
share more elongate valvae, with centrad located dorsal process, nar- 
row terminus, and thin terminal hook (Fig. 2). Epargyreus windi ex- 
hibits a large, but centrally limited, silver-white patch on the hindwing 
under-surface, which might further suggest affinity to E. spanna. 

The above genitalic similarities do not necessarily override the close 
sister species relation posited by some for E. spanna, E. antaeus and 
E. zestos. No one has done a cladistic analysis of Epargyreus, but the 
apparent affinity of genitalic and wing characters summarized above 
suggests that Antillean Epargyreus species exhibit character states root- 
ing them as primitively in phylogenetic diagrams as their mainland 
congeners. The suggested genitalic affinities of E. spanna are remark- 
able because they suggest phylogenetic and zoogeographic relations 
contrary to the common practice of associating Antillean endemics as 
subspecies of various mainland or Greater Antillean taxa (Riley 1975, 
Clench 1965). If the latter were true, one would expect the wing char- 
acter affinities for E. spanna, E. antaeus, and E. zestos to be closely 
supported by characters of the genitalia. 

Johnson and Matusik (1986) noted a similar situation in the genitalic 
characters of a new, apparently endemic Hispaniolan Tmolus (Ly- 
caenidae). This species does not resemble its most geographically prox- 
imate Antillean or mainland congener. Steven Steinhauser [Allyn Mu- 
seum of Entomology] (pers. comm.) concurs regarding the genitalic 
characters of Astraptes christyi Sharpe of Hispaniola. Astraptes chris- 
tyi has long been treated as a subspecies A. xagua Lucas (Riley 1975), 
even though it differs from that taxon in characters of the wing. Gen- 
italic examination of A. christyi indicates species status and affinities 
not simply reducible to sympatric E. xagua. Schwartz and Miller (1985), 
in describing a new endemic Hispaniolan Strymon, demonstrate other 
affinities than might be presupposed from the most geographically 
proximate congener. We recently collected in Hispaniola an unde- 
scribed species of Nesiostrymon which cannot be regarded as N. celida 
aibinito Comstock & Huntington of Hispaniola, or of the N. celida 
Lucas complex now divided into island subspecies. The practice of 
placing Antillean populations of butterflies as subspecies of other, more 
common Antillean or mainland species seems to have resulted from 
certain zoogeographic assumptions. Many authors believed that Antil- 
lean butterfly distributions represent results of recent (Pleistocene and 
post-Pleistocene) waif dispersal (Comstock & Huntington 1943, Clench 
1965). Such assumptions have been strengthened by limiting compar- 
isons to wing patterns and interpreting differences in these as “‘varia- 
tion’”’ without reference to structural characters. 


VOLUME 40, NUMBER 1 63 


In zoogeography there is currently an increased appreciation of the 
possible correlation of late Mesozoic and Cretaceous plate tectonic 
splitting, and allopatric speciation of biological populations now rep- 
resenting various areas of endemism (Rosen 1975). This view predicts 
Antillean endemics may exhibit arrays of characters as primitive as 
any of their mainland counterparts. It is apparent that the affinities of 
E. spanna do not support the simplistic island subspecies view of Ca- 
ribbean biogeography. We expect that future cladistic analyses of 
Epargyreus and other butterfly groups will suggest the early origin of 
many endemic Antillean taxa. 


ACKNOWLEDGMENTS 


We thank Albert Schwartz (Miami Dade County Community College, Miami, Florida) 
and Lee D. and Jacqueline Y. Miller (Allyn Museum of Entomology, Florida State Mu- 
seum, Sarasota, Florida) for helpful information. Albert Schwartz and H. A. Freeman 
(Garland, Texas) reviewed the manuscript and made useful suggestions. Frederick H. 
Rindge (AMNH) kindly facilitated access to the AMNH collections, and also commented 
on the manuscript. 


LITERATURE CITED 


BROWN, F. M & B. HEINEMAN. 1972. Jamaica and its butterflies. E. W. Classey, Ltd., 
London. 478 pp. 

CLENCH, H. K. 1965. A synopsis of the West Indian Lycaenidae with remarks on their 
zoogeography. J. Res. Lepid. 2:247-270. 

CoMSTOCK, W. P & E. I. HUNTINGTON. 1943. Lycaenidae of the Antilles (Lepidoptera: 
Rhopalocera). Ann. New York Acad. Sci. 45:119-180. 

Evans, W. H. 1952. A catalogue of the American Hesperiidae. II. Pyrginae. British 
Museum (Natural History), London. 246 pp. 

Gaul, F. & A. SCHWARTZ. 1988. The second specimen of Epargyreus spanna (Hesper- 
iidae). J. Lepid. Soc. 37:170-171. 

GLoyp, L. K. 1982. The original definition and purpose of the term allotype. Syst. 
Zool. 31:334-336. 

FREEMAN, H. A. 1969. Records, new species, and a new genus of Hesperiidae from 

- Mexico. J. Lepid. Soc. (suppl. 2). 62 pp. 

1977. Six new species of Hesperiidae from Mexico. J. Lepid. Soc. 31:89-99. 

FRIZZELL, D. L. 1988. Terminology of types. Am. Midl. Nat. 14:637-668. 

JOHNSON, K. & D. Matusik. 1986. A new species of Tmolus (Lycaenidae) from His- 
paniola. Addendum in A. Schwartz, Butterflies of Hispaniola, Mus. Hist. Nat., Re- 
publica Dominicana (in press). 

RILEY, N. D. 1975. Field guide to the butterflies of the West Indies. Collins, London. 
244 pp. 

ROSEN, D. E. 1975. A vicariance model of Caribbean biogeography. Syst. Zool. 24:431- 
464. 

SCHWARTZ, A. & J. Y. MILLER. 1985. A new species of Strymon (Lycaenidae) from 
Hispaniola. Bull. Allyn Mus. 99:1-6. 

SMITH, H. M. 1988. More on allotypes. Syst. Zool. 32:454—455. 





GENERAL NOTES 


Journal of the Lepidopterists’ Society 
40(1), 1986, 64-65 


EMERGENCE OF ADULT ECTOMYELOIS MURISCUS (DYAR) 
(PYRALIDAE) FROM A POD OF THEOBROMA SIMIARUM 
DONN. SMITH (STERULIACEAE) IN COSTA RICA 


The pyralid moth Ectomyelois muriscus (Dyar), a species widely distributed in Central 
America, northern South America, and the West Indian archipelago, undergoes its life 
cycle in the pods of Theobroma cacao Linnaeus (Sterculiaceae) and other fruits (Heinrich 
1956, American moths of the subfamily Phytcitinae, U.S. Natl. Mus. Bull. No. 207). I 
have been unable to locate published records of this moth species infesting pods of other 
Theobroma, a genus represented by several species in tropical America (Cuatrecasas 
1964, Cacao and its allies—A taxonomic revision of the genus Theobroma, Contrib. U.S. 
Natl. Mus. 35:379-614). Given the marked differences in external texture, pubescence, 
and other morphological features of pod walls among Theobroma species, one might 
expect some degree of ovipositional selectivity to exist for moth species associated as 
larvae with pods of these neotropical tree species. Here I report the emergence of 41 
adults of Ectomyelois muriscus from one rotted and dried pod of Theobroma simiarum 
Donn. Smith in Costa Rica, representing a new host record for the moth, and for a 
Theobroma species with mature pods differing markedly in pod-wall texture from the 
previously reported host, T. cacao. 

One of 12 fallen, mature, and decaying pods of T. simiarum was collected beneath 
one of four trees of this species in the “Theobroma and Herrania garden” on the grounds 
of the “Centro Agronomico Tropical de Investigaciones y Ensenanza’’ (CATIE) at Tur- 
rialba (9°55’'N, 83°41'W; about 600 m elev.), Cartago Province, Costa Rica in mid-August 
1984. The 26 x 8 cm brown pod had no external insect emergence holes at the time it 
was collected. Subsequently the pod was kept on a desk in an office. Following an initial 
emergence of a few moths, I confined the pod in a plastic bag in the office. All adults 
were kept, and voucher specimens sent to the U.S. Dep. Agr. Systematic Entomology 
Laboratory (U.S. National Museum) for determination. 

Between 20 October and 7 December 1984, 42 moths emerged from the pod, with an 
approximately 1:1 sex ratio. From one to four moths emerged on a given day during this 
period, but there were many days when no moths emerged. Most moths emerged before 
0800 h. Several freshly-eclosed moths clung motionless to the pod for several hours, and 
flew only when disturbed. By the time the last moth emerged, only three exit holes were 
found on the external surface of the pod. Clearly, several moths used the same exit holes 
for emergence. Each exit hole had a 10-25 mm long silken tube externally, apparently 
built by larvae inside the pod wall and pushed out at the time of multiple eclosions. But 
eclosion behavior was not observed. Nor did I open the pod to determine where larvae 
were feeding, as the intact fruit was necessary for other research purposes. 

The adults exhibited a staggered emergence pattern, because the emergence period 
lasted about six weeks. The female moth probably deposits clusters of eggs on the external 
surface of the sand-papery-rough pubescent pod, since adults appeared to emerge in 
clusters from a few exit holes. Perhaps this particular pod received three different egg 
batches. Assuming the observed laboratory emergence pattern was similar to that occur- 
ring in nature, a brood of E. muriscus emerges near the end of the Turrialba rainy 
season, and before the short, erratic dry season. The availability of decaying pods of T. 
simiarum varies greatly throughout the year, suggesting a changing pod supply for pod 
herbivores or pod saprotrophs (whichever the case may be). 

Given the previously reported association of E. muriscus with T. cacao in both Central 
and South America, my discovery of this moth species in a pod of T. simiarum may not 
be surprising. Theobroma simiarum is one of two species of the genus endemic to 
Costa Rica. Given the broad geographical range of Ectomyelois muriscus in tropical 
America, it undoubtedly has other natural hosts, possibly species of Theobroma other 


VOLUME 40, NUMBER 1 65 


than cacao or simiarum. The dense, thick tomentum (pod wall external surface) may 
represent a suitable oviposition substrate for Ectomyelois muriscus, but other surface 
textures must also be suitable given the marked difference in this feature between Theo- 
broma cacao and T. simiarum. Larvae of Ectomyelois muriscus most likely tunnel 
through the woody epicarp and softer mesocarp tissues of the pod. Yet they may infest 
pods once the latter are into advanced stages of decay, perhaps rendering pod-wall tissues 
more penetrable to larvae. 

Near the end of the rainy season at this locality, mature pods of various species of 
Theobroma are available, in addition to those of T. cacao, the most abundant species 
due to large commercial plantations. When the dry season arrives near the end of De- 
cember, dryness may trigger a large moth emergence, a pattern somewhat different than 
that observed in the office. The very dry conditions of the office may have mimicked 
the dry season for moth larvae and pupae present inside the T. simiarum pod, leading 
to a staggered emergence as conditions became increasingly dry. 

This research was funded by the American Cocoa Research Institute of The United 
States of America. I thank D. C. Ferguson for determining the moth and providing the 
Heinrich reference. The technical assistance of Susan Sullivan Borkin is appreciated. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 65-66 


THE FEMALE OF PAPILIO XANTHOPLEURA 
GODMAN & SALVIN (PAPILIONIDAE) 


Before 1985, literature concerning Papilio xanthopleura Godman & Salvin stated that 
its female occurs in two forms: a “normal” female resembling the male, and a large 
yellow one, form diaphora Staudinger (Staudinger 1891, Deut. Entomol. Z. [Iris] Lepid. 
4:61-158; Rothschild & Jordan 1906, Novit. Zool. 13:412-752; Jordan 1907, in Seitz, 
Macrolepidoptera of the World, Vol. 5, Alfred Kernen Verlag, Stuttgart, 592 pp.; Munroe 
1961, Can. Entomol. Suppl. 17, 51 pp.; D’Almeida 1965, Catalogo dos Papilionidae 
Americanos, Soc. Braz. Entomol., Sao Paulo, 366 pp.; D’Abrera, Butterflies of the Neo- 
tropical Region, Part 1, Papilionidae and Pieridae, Lansdowne Editions, East Melbourne, 
172 pp.). None of the literature illustrates a xanthopleura female. 

Johnson, Rozycki and Matusik (1985, J. N.Y. Entomol. Soc. 93:99-109), examined the 
type and other specimens of diaphora, and showed that the type and all known repre- 
sentatives of diaphora are males, and male genital and wing characters in diaphora 
indicate it is not conspecific with xanthopleura. As a result, diaphora was accorded 
species status, it became apparent that females of diaphora are presently unknown in 
collections, and no “normal” females of xanthopleura were in the following major col- 
lections: Allyn Museum of Entomology, American Museum of Natural History (AMNH), 
British Museum (Natural History), Carnegie Museum of Natural History, Collection of 
David Matusik (Skokie, Illinois), Collection Dep. de Zoologia, Universidade Federal do 
Parana (Brazil), Collection of Ernesto W. Schmidt-Mumm (Bogota, Colombia), Collection 
of Rick Rozycki (Chicago, Illinois), Collection Tommasso Racheli (Rome, Italy), Instituto 
de Zoologia Argricola Maracay (Venezuela), Museu Nacional, Rio de Janeiro (Brazil), 
Museo de Historia Natural “Javier Prado” (Lima, Peru), National Museum of Natural 
History (Smithsonian Institution), and the collection of a commercial dealer noted for 
his holdings in unusual Papilionidae. 

Therefore, we borrowed a female of xanthopleura (Fig. 1A, C) from the Staudinger 
Collection (Zoologisches Museum der Humboldt Universitat, Berlin, German Democratic 
Republic [ZMH]). The female resembles male xanthopleura on the wing undersurface 
but, contrary to the above literature, differs markedly from the male on the upper surface 
of the wings. Males of xanthopleura are black above except for brilliant “powder green’ 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Papilio xanthopleura and P. diaphora, with forewing length (base to apex) 
in parentheses, B—D, upper surfaces of wings, to same scale, A, undersurface, to different 
scale. A, P. xanthopleura female (67.0 mm), Iquitos, Peru, ZMH; B, P. diaphora type 
male (71.0 mm), Manicoré, Brazil, ZMH; C, P. xanthopleura female (of 1A); D, P. 
xanthopleura male (57.0 mm), Campana [sic], Brazil, AMNH. 


in the vein interspaces of the hindwing (Fig. 1D); females are powder green over the 
entire upper surface of both wings (Fig. 1C). Female xanthopleura are larger than male 
xanthopleura, but neither exceeds the large size of male diaphora. As noted by Johnson, 
Rozycki and Matusik, the mean single forewing length (base to apex) of known diaphora 
males exceeds that of examined xanthopleura males by 12.8 mm and the examined 
xanthopleura female by 3.0 mm. Thus, wing character differences in the genders of 
these taxa vary far more than the literature has indicated. 

We thank Prof. H. J. Hannemann for loan of the diaphora type and various xantho- 
pleura specimens. Phillip Ackery, K. S. Brown, O. H. H. Mielke, L. D. Miller, John 
Rawlins, R. K. Robbins, Tommasso Racheli, and E. W. Schmidt-Mumm aided in survey- 
ing various collections. 


KURT JOHNSON, Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024; Rick ROZYCKI, 5830 
South McVicker Avenue, Chicago, Illinois 60638; AND DAvID MATUSIK, Department of 
Entomology, Field Museum of Natural History, Roosevelt Road at Lake Shore Drive, 
Chicago, Illinois 60605. 


VOLUME 40, NUMBER 1 67 


Journal of the Lepidopterists’ Society 
40(1), 1986, 67 


PLACEMENT AND FATE OF MONARCH BUTTERFLY PUPAE 
IN NORTHERN CALIFORNIA 


The placement on a substrate and the subsequent fate of Danaus plexippus (L.) (Dana- 
idae) pupae were determined on a roadside strip at two sites SE of Davis, Yolo Co., 
California, in the Sacramento Valley from mid-July to October 1966. 

The first site was 3 km south of Davis. It was a strip 4 m wide and about 400 m long 
containing several clumps of Asclepias fascicularis, other green plants, and dry grass 
between a paved road and a woven wire fence. The second site was similar and 6 km 
SSE of Davis. Both were in flat, mostly agricultural land, and near alfalfa, sugar beets, 
and grains. The only trees were around a dwelling and an old church and cemetery. 

Each site was thoroughly searched on weekends and occasionally during the week 
from 14 July to 30 October 1966. Each pupa was marked with a paper tag located nearby 
with a serial number and date of discovery. 

Vertical distributions of pupae were from ground level to 1 m. One pupa was found 
4 m high on an old creosoted pole. 

Choices of substrate for 409 pupae were: A. fascicularis, 17.7%; other green plants, 
6.2%; dry grass, 47.7%; wire fence, 25.5%; and wooden fence posts, 2.9%. Only 4% of 
the pupae on the A. fascicularis were placed after 31 August. On the wire fence, 6% 
were placed in July, 25% were placed in August, 53% in September, and 16% in October. 
On dry grass, 24% were placed in August, 53% in September, and 23% in October. Pupae 
are rarely placed on Asclepias spp. in southeastern Canada (Urquhart 1960, The Monarch 
Butterfly, Univ. Toronto Press, 361 pp.). 

Following initial observations, several categories of pupal fate were defined but only 
three are reported (Table 1). The latter are: discolored and dead, empty shell left after 
eclosion, and disappeared leaving no evidence of fate (because pupae were marked with 
white tags, passersby may have taken some, but predation seems more likely). 


TABLE 1. Danaus plexippus pupal fate by substrate, Davis, California, 1966. 


Number of pupae 


Asclepias Other green 
Fate Dry grass fascicularis plants Wire fence Total 
Died 83 25 9 21 138 
Eclosed 74 32 14 35 155 
Disappeared 54 18 4 23 99 
Total 211 75 27 79 392 


Without regard to substrate’ 39.4% eclosed, 35.1% died, and 25.5% disappeared (Table 
1). Of the 99 that disappeared, 55% were on dry grass, 28% on the wire fence, and 24% 
on green plants. The data were analyzed in a standard contingency table; x? = 7.18, df = 
6, .500 > P > .250. The sample may be too small, but tentatively, pupae on green plants 
appear to be safest from predators. A higher percentage of pupae eclosed on green plants 
than on dry grass or the wire fence. 

I thank A. M. Shapiro of the University of California, Davis, for reviewing the manu- 
script and doing the statistical test. 


LESLIE V. SMITH, 7589 Twin Oaks Ave., Citrus Heights, California 95610. 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Journal of the Lepidopterists’ Society 
40(1), 1986, 68 


ABERRANT POSTMEDIAL SPOTS IN 
ICARICIA ICARIOIDES LYCEA 
AND I. ACMON (LYCAENIDAE) 


Several nearctic lycaenids tend to abnormal expression of ventral postmedial spots. 
This generically takes the form of some or all spots being elongated. Illustrations recently 
published include a multiple occurrence in Satyrium calanus falacer (Godart) (Wright 
1981, J. Lepid. Soc. 35:158-159). Single occurrences have been shown for Satyriwm 
acadica acadica (W. H. Edwards) (Leeuw 1979, J. Lepid. Soc. 33:204—205), Glaucopsyche 
lygdamus couperi Grote (Neil, 1983, J. Lepid. Soc. 37:258), Euphilotes rita rita (Barnes 
& McDunnough), Icaricia acmon texana Goodpasture (Holland 1980, J. Res. Lepid. 19: 
88-95) and Icaricia acmon lutzi (dos Passos) (Cannon, J. Lepid. Soc. 39:329-330). The 
similarity of the two aberrant acmon specimens is striking, especially since the acmon 
lutzi is from Idaho and the acmon texana is from New Mexico. 

To this list of aberrant lycaenids, I now add Icaricia icarioides lycea (W. H. Edwards). 
Figure 1 shows an aberrant, an intermediate and a typical specimen. About 5% of 100 
specimens were found to have noticeably enlarged postmedial spots. The two atypical 
examples illustrated here represent the most extreme. 

I call attention to a well illustrated and useful article on the mechanics of color pattern 
formation and malformation in butterflies and moths (Nijhaut 1981, Sci. Am. 245(5): 
140-151). This work seems not to have received the recognition it should among lepi- 
dopterists. 


RICHARD HOLLAND, 1625 Roma NE, Albuquerque, New Mexico 87106. 





Fic. 1. Icaricia icarioides lycea ventral surface. Left, aberrant male, Corral Tank, 
Canyon de Los Corrales, NE slope, Jemez Mts., Rio Arriba Co., NM, 27.V.84, 2,438 m; 
Middle, intermediate male, 1.6 km N of Poncha Pass, Chaffee Co., Colo., 5. VII.63, 2,743 
m; Right, normal male, 20.9 km W of Espanola on road to Santa Clara Peak, E slope, 
Jemez Mts., Rio Arriba Co., NM, 18.V1.83, 2,499 m; all leg. R. Holland. 


VOLUME 40, NUMBER 1 69 


Journal of the Lepidopterists’ Society 
40(1), 1986, 69-71 


NOTES ON A COSTA RICAN “MONKEY SLUG” (LIMACODIDAE) 


Little is known about the life cycles, larval food plants and other aspects of natural 
history of neotropical limacodids (Dyar 1924, Limacodidae, in Macrolepidoptera of the 
World, Vol. 6. American Heterocera [A. Seitz, ed.], A. Kernan Verlag, Stuttgart). Herein 
I describe the final instar caterpillar, some aspects of caterpillar behavior, and one larval 
food plant for Phobetron hipparchia Cramer in northeastern Costa Rica. Dyar mentions 
that P. hipparchia caterpillars feed on “different forest trees” and that this species occurs 
in Mexico, Panama, Ecuador, Colombia, Venezuela, Guiana, Brazil, and Argentina. 

On 27 February 1985, two late-instar caterpillars of P. hipparchia were collected from 
one 6 m tall Gliricidia sepium (Jacq.) Steud. (Papilionoideae: Galegeae, Robiniinae) 
supporting vanilla vines at “Finca La Tirimbina,” near La Virgen, Sarapiqui District, 
Heredia Province (10°23’N, 84°07’W;; 220 m elev.). The tree was one of several thousand 
G. sepium planted there for vanilla production (Allen & Allen 1981, The Leguminosae: 
A source book of characteristics, uses and nodulation, Univ. of Wisconsin Press, Madison, 
Wisconsin, 812 pp.). The caterpillars were placed in a clear-plastic bag, along with 
cuttings of G. sepium, for rearing to adulthood. 

A thorough search of the G. sepium having the caterpillars revealed no other individ- 
uals of P. hipparchia, as was also the case for an additional eight trees of this species 
examined in the same area. Both caterpillars were discovered on the dorsal (upper) 
exposed surfaces of old, tough leaves (Fig. 1), and about 1 m apart at eye level (about 
1.8 m above the ground). At the time of discovery, the majority of G. sepium trees at 
the site were without flushes of new (fresh) leaves. From a distance of about 1 m, the 
caterpillars resembled curled, dry leaves (Fig. 1). 

Within three days after collection, both caterpillars molted to the final instar, exhibiting 
little change in overall appearance from the previous instar. The final instar lasted only 
a few days; both caterpillars formed loose silken cocoons in the leaves by 7 March. Each 
cocoon (Fig. 1) consisted of a thin sheet of silken mat across several leaves, and dorsally 
mostly the larval tubercles shed during spinning. The pupal stage lasted about one month 
(under laboratory conditions of 65—70°C and 30-40% RH). Both adults emerged between 
1500-1600 h. One was female, the other a male (Fig. 1). 

At the time of discovery, the caterpillars were both about 24 mm long and 20 mm 
wide, including the greatest expanse of the laterally positioned “horns” (Fig. 1). When 
the caterpillar was motionless on a leaf, the horns were held laterally against the leaf 
surface (Fig. 1). When moving, the caterpillar appeared to be rocking back and forth, 
with a slight rotation of the horns. The squarish, angular body profile of a motionless 
caterpillar on a leaf (Fig. 1) became spherelike when the caterpillar was disturbed: the 
caterpillar curled itself into a ball, and partially tucked in and interlocked some of the 
lateral horns. Several sustained prods with a forceps were needed to ellicit this curling 
behavior. 

The overall color of the caterpillar consisted of a patchwork of brown shades. Cramer 
(1791, Uitlandsche Rupsen, Supplement) reported the larva (Plate XVIII) of P. hippar- 
chia to be light-brown in color. As in all Limacodidae, the head capsule (about 4 mm 
diam) was small and hidden at all times. Both the head capsule and thoracic legs were 
glossy orange. The first two thoracic segments were almost translucent, and without 
prominent tufts of setae or lateral extensions of the cuticle. The third thoracic segment 
had a ringlet of six bulbous, orange tufts of hairs. The lateral horns were present on the 
first three abdominal segments (one pair per segment), and dorsally were darker brown 
than below. The horns of the third segment were dorsally more markedly brown than 
those of the previous two segments. Those of the fifth abdominal segment were dark 
brown dorsally, while those of segments 7-9 were light brown (tan) dorsally. No lateral 
horns were present on abdominal segments 4 and 6. The lateral horns appeared to be 
extensions of the cuticle, and were covered with short setae (see Dyar 1896, J. New York 
Entomol. Soc. 4:167-190). Dorsally, each abdominal segment had a dark brown rectan- 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 1. Phobetron hipparchia. Left top and bottom: Final-instar caterpillars on leaves 
of Gliricidia sepium. Right top: Cocoon. Right bottom: Reared male (above) and female 
(below). 


gular patch, with a light spot in the center and even darker borders (Fig. 1). A dorsal- 
medial raised line of tan-colored setae ran lengthwise on both thoracic and abdominal 
regions. The body of the caterpillar was 8 m wide at the thickest point within a day of 
cocoon formation. Following cocoon formation there was no noticeable change in the 
colors of the larval cuticle, even though it became part of the cocoon. 


VOLUME 40, NUMBER 1 ral 


Phobetron hipparchia, the single species of the genus represented in the neotropical 
region, apparently gets the name “monkey slug” from the curiously shaped caterpillar 
stage. One of the two North American species of the genus, P. pithecium (J. E. Sm.) is 
the “hag moth,” and its caterpillars feed on a broad range of trees, none of which 
apparently is within the Leguminosae (Papilionoideae) (Covell 1984, A field guide to the 
moths of eastern North America, Houghton Mifflin Co., Boston, 496 pp.). Dyar (1896, 
op. cit.) noted that P. pithecium larvae perch on the undersides of leaves until the last 
instar, and that Phobetron larvae in general cryptically resemble dead leaves. Even 
though perched on the upper sides of leaves, the final-instar larva of P. hipparchia 
appears cryptic, resembling the yellow, brown, and green blotch pattern of older G. 
sepium leaves. The observed pattern of cocoon construction, in which cast-off tubercles 
are added to the silk during spinning, is considered typical by Dyar for New World 
Phobetron, enhancing crypsis of the pupal stage. 

Of the several genera and species of North American Limacodidae discussed in Covell, 
none apparently utilize legumes as larval food plants. Yet an outstanding feature of these 
moths in general appears to be their highly polyphagous food habits as caterpillars (Dyar 
1924, op. cit.; Covell, op. cit.). Several genera and species of Limacodidae feed on 
legumes in Australia. 

McFarland (1979, J. Lepid. Soc. 33: Supplement, 72 pp.) reports that Australian li- 
macoids are associated with Acacia and other legume genera, and that caterpillars of 
some species invariably occur on old, tough leaves and stems of food plants as in the 
present observations. Within the neotropical region, the limacodid Sibine apicalis (Dyar), 
or “gusano montura,’ sometimes defoliates banana (Musa spp.) trees (Jaramillo & Ji- 
menez 1974, Turrialba 24:106-107). Thus both dicotyledenous and monocotyledenous 
larval food plants for the Limacodidae are known from the neotropical region. Cramer 
reported P. hipparchia on Granadilla (Passifloraceae). Eucalyptus spp. (Myrtaceae) are 
major larval food plants for the Limacodidae in Australia and Africa (McFarland; Se- 
vastopulo 1983, J. Lepid. Soc. 37:91, respectively). But in East Africa some limacodids 
feed on indigenous legumes (Acacia spp.), but not introduced species of the family. 
Gliricidia is endemic to the neotropical region, and given the great evolutionary diver- 
sification of the papilionoid legumes in tropical America (Richards 1964, The Tropical 
Rain Forest, The Univ. Press, Cambridge, England, 437 pp.), one might expect to dis- 
cover several other legume larval food plants for P. hipparchia. Gliricidia is widely 
distributed throughout the American tropics, both in natural habitats, as a result of its 
extensive use as a shade tree for cacao and coffee, and as a support for vanilla vines. 

In spite of several years of casual observation during both dry and rainy seasons, I did 
not notice other P. hipparchia caterpillars on the trees. 

This research was a by-product of a grant from The American Cocoa Research Institute 
of The United States of America. I thank J. Robert Hunter for allowing me access to 
Finca La Tirimbina. Adult moths reared in this study have been deposited in the collec- 
tions of the Milwaukee Public Museum. Detailed comments of the reviewers were most 
helpful, and one reviewer provided me with the Cramer (1791) and Dyar (1896) refer- 
ences, for which I am grateful. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53238. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 72-73 


BOOK REVIEWS 


MILKWEED BUTTERFLIES, by P. R. Ackery and R. I. Vane-Wright. 1984. British Museum 
(Natural History), Cromwell Road, London SW7 5BD, England. v-ix + 425 pp., includ- 
ing 261 figures, 12 colored and 73 halftone plates, quarto size. Price £ 50. Also published 
in U.S. with Cornell Univ. Press, Ithaca, New York. 


The full title of this book, “Milkweed butterflies: their cladistics and biology, being an 
account of the natural history of the Danainae, a subfamily of the Lepidoptera, Nym- 
phalidae” states the goals of Ackery and Vane-Wright’s study. They succeed admirably 
in this difficult task. While the major thrust of the book is a taxonomic revision of the 
danaines, there is a wealth of well documented biological information about distribution, 
behavior, life history, chemical attraction and defense, genetics, ecology, mimicry and 
faunistics. This makes the book not only valuable to the systematist, but also to the worker 
in other biological disciplines. 

The fact that the revisionary section is a cladistic study probably will bother phenet- 
icists and at least some evolutionary taxonomists, but this does not concern the authors. 
It is reassuring to see a study that is done strictly on characters, and draws no systematic 
conclusions not supported by those data; the authors freely admit that certain relations 
are not shown by the characteristics they used, and suggest that these problems may be 
solved later by the addition of new characters. What is speculation by the authors is 
clearly so labeled, and definitive statements are not made without reference to underlying 
reasons. The revision is, therefore, highly scientific; and it is a pleasure to have the authors 
lead one through the reasoning to their conclusions without the appearance of the occult 
that one is so often left with in systematic revisions where conclusions are “correct” 
simply because authors state they are. 

The cladistic section of the book involves discussions of the characters treated and the 
classification scheme derived from them. Each character is discussed, numbered in the 
text and, perhaps most importantly, is illustrated clearly in the figures, along with alter- 
nate character states. Again, one can follow the reasoning through a logical progression 
to the classification adopted. There are some valuable, but unconventional, thoughts on 
classification intertwined with the data, such as those on clades and polytypic species 
(pp. 20-22), including the concepts of “‘cladospecies” and “paraspecies”’, and those im- 
mediately following on larval versus adult characters. Their suggestions for further sys- 
tematic research on the danaines (pp. 61-66), most of which the authors promise to 
tackle later are valuable and thought-provoking. The book would have been even more 
useful had it considered the myriad subspecies, but an attempt to analyze cladistically 
some 2,000 names would have been humanly impossible, and the result would not have 
been economically feasible to print. Nevertheless, the handling of the subject is reasonable 
enough to make a confirmed cladist of the reader: if only people of that taxonomic school 
were always so rigorously tied to logic! 

The authors anticipated that the revision might be controversial because it upsets 
prevailing nomenclature. Ackery and Vane-Wright comment about the genus “Danaus” 
of authors. “Danaus” is at best a grade taxon and is probably polyphyletic, perhaps 
paraphyletic. A number of species are placed in genera far removed from their “con- 
ventional” placement, but the authors recognize the difficulty that others might have 
accepting these new assignments, stating (p. 8), “. . . because of the still overriding 
influence of ‘Seitz’ and the acceptance of ‘Danaus’, we can be sure that a dual nomen- 
clature, with ‘Danaus’ sita for Parantica sita, ‘Danaus’ similis for Ideopsis similis, 
‘Danaus’ hamata for Tirumala hamata and so on, will unfortunately continue in exis- 
tence for a considerable length of time—probably until about A.D. 2179 judging by 
past performances!” Such a statement might be made about any revision that accepts a 
different nomenclature than that in a “standard” work, but Ackery and Vane-Wright 
present a large and impressive body of data, and it is up to their detractors (if any) to 
examine such data in detail and show where they think the authors are wrong. I hope 


Ackery and Vane-Wright are incorrect on the period of time until their nomenclature 
is accepted. 


VOLUME 40, NUMBER 1 73 


Ackery and Vane-Wright have gathered an impressive array of biological data on the 
danaines which are summarized in Part 2: Biology (pp. 67-102). They have also gener- 
ated a large body of data on co-mimicry and faunistics (pp. 1038-158). Both of these 
sections should be of great interest to ecologists, evolutionists and other nontaxonomists, 
as well as to systematists in the broadest sense. Part 4 consists of identification keys, often 
utilizing novel characteristics not stressed in the typical key; and all data are summarized 
in Part 5, the specific taxonomic and biological catalogue (pp. 173-245), an impressive 
compendium of information that should convince even the most skeptical. There are 
some new synonyms, combinations and taxa, but finding them requires some searching 
because they are buried throughout the text. Short of a section summarizing changes, 
there is no other way the data could have been presented conveniently. A short adden- 
dum follows, and precedes an exhaustive bibliography of 86 pages. 

The work is remarkably free of typographical errors—the one I noticed was the ren- 
dering of Japan as “Japen” on p. 21. It is an attractive book and easy to follow. The 
illustrations are uniformly of high quality, including line drawings, colored plates, half- 
tone plates, and 394 additional figures illustrating all aspects of danaines and their biol- 
ogy. Colored plates depict some danaine mimetic associations in the Philippine and 
Indonesian Islands. It is too bad that illustrations are not cross-referenced to the pages 
on which descriptions occur (the pages with descriptions do have references to plates), 
but this is a minor complaint; also each species is mentioned several times in the text, 
and to which page would the plate reference refer? The comprehensive Index clearly 
leads the reader to any place a taxon is discussed. 

I must consider this work to be one of the major taxonomic revisions of this century, 
and it is a good book for the reader who is not taxonomically inclined. The authors have 
attacked a problem, solved much of it, and have honestly admitted those parts that have 
resisted solution so far. To say that I am impressed with this work is an understatement; 
it is the kind of work that one always hopes to be able to do. The book, even though 
expensive, is well worth the cost; I would recommend it to anyone interested in well 
explained and defended cladistic analyses, in systematics of Lepidoptera, or in the Danai- 
nae. 


LEE D. MILLER, Allyn Museum of Entomology of the Florida State Museum, 3621 
Bay Shore Road, Sarasota, Florida 33580. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 74 


A MONOGRAPH OF THE BIRDWING BUTTERFLIES. Volume I, parts 1-8, the subgenera 
Aetheoptera, Ornithoptera, Schoenbergia; Volume II, parts 1-2, the genera Trogonop- 
tera, Ripponia, Troides (partim), by J. Haugum and A. M. Low. Vol. I. 308 pp., 12 
plates. Vol. II. 240 pp., 12 plates. Scandinavian Science Press, Ltd., Klampenborg, Den- 
mark. 1978-1984. 


No other group of butterflies has attracted as much attention or interest as the birdwing 
butterflies. In this two-volume set, the authors provide the most detailed analysis of these 
showy butterflies to date. The first part of Vol. I, for example, on Aetheoptera, deals 
with only two species, yet numbers 84 pages; and Vol. II, part 1, which describes three 
species, numbers 104 pages. The text of this work will complement the beautiful illus- 
trations in the recently published Birdwing Butterflies of the World, by B. D’Abrera. 

Volume I covers the genus Ornithoptera, Volume I, Trogonoptera, Ripponia, and 
Troides. The genera and species are introduced with descriptions and notes on biology, 
followed by keys. However, no keys are given for most subspecies, which means that 
most (as with so many subspecies of butterflies) must be determined by geographic 
locality. Each species and subspecies is described in detail, and notes are given on the 
phylogenies of each taxon. Several illustrations, including distribution, accompany the 
description of each taxon. The authors tend to recognize almost every described taxon, 
and include descriptions of several “forms” or aberrations. Even though the authors 
recognize that formal names given to such infraspecific forms are not nomenclatorally 
valid, I find it annoying to see several new names applied to these forms. 

A group as well known and as popular as the birdwing butterflies is bound to generate 
controversy in the literature, and such is the case here. I found the authors to be partic- 
ularly critical of D’Abrera, often disagreeing with what is said in his volume. For ex- 
ample, D’Abrera recognizes only two subspecies of Ornithoptera goliath, Haugum and 
Low, five. D’Abrera considers O. richmondia and O. urvillianus separate species from 
O. priamus, not so Haugum and Low. Ripponia is used for hypolitus, but D’Abrera 
considers it a Troides. 

For all the detailed analysis given for each subspecies, I find it disappointing that no 
quantitative data were used to back up the authors’ assertions that these taxa are, indeed, 
taxonomically distinct. Most of the diagnostic comments are qualitative, as two examples 
will show: Males of O. tithonus “waigeuensis may be recognized by having a narrow 
HW [hind wing] with a notable reduction of the apical area; the wing is even further 
modified and less angular at the apex than in subspecies misresiana and tithonus ...,” 
and “The male HW [of T. amphrysus ruficollis f. loc. euthydemus] tends to be more 
rounded than in ruficollis on average, also the FW [fore wing] may be broader ... ,” 
which means to the reader that geographic locality will still be the only way to identify 
subspecies. 

Puzzling also is the uneven treatment of “Material examined.” For some species, a 
detailed listing is given, for others, the data are incomplete, as for O. goliath atlas: 
“photographs of a further series of E. Weyland Mts. imagines” [how many?]. The loca- 
tions of types are, unfortunately, not addressed for all taxa. No cladistic analysis was 
carried out on this group, although I hope the authors may consider this in their final 
volume. 

The books are handsomely bound, the printing, layout, and illustrations are good, but, 
alas, errors in typography and syntax abound. The color photos of Vol. I are excellent, 
but those of Vol. II have a white halo within the marginal black areas of the wings 
figured. 

Despite their shortcomings, these volumes represent the best compilation of data for 
these magnificent butterflies. As one who is fascinated by these gorgeous insects, I heartily 
recommend this work for every lepidopterist interested in Old World Papilionidae. 


ROSSER W. GARRISON, 1030 Fondale St., Azusa, California 91702. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 75-76 


OBITUARIES 


ARTHUR C. ALLYN (1913-1985) 


Dr. Arthur Cecil Allyn, life member of the Lepidopterists’ Society and Director Emer- 
itus of the Allyn Museum of Entomology, died on 22 March 1985, after a lingering 
illness. He will be remembered by the lepidopterological community for his generosity, 
dedication, and service to the science. 

Dr. Allyn was born in Evanston, Illinois on 24 December 1918, completed primary 
and secondary schooling there, and attended Dartmouth and Beloit Colleges. He received 
a D.Sc. from the University of Florida in 1981 in recognition of his accomplishments in 
and service to entomology. He is survived by his wife, Dorothy D. Allyn, and three 
children, David D. Allyn, William N. Allyn and Dorothy A. Lavick, as well as eight 
grandchildren. 

Dr. Allyn had a successful and diversified business career with international interests 
in oil, maufacturing, farming and sports in many countries, including Australia, Indo- 
nesia, Canada, South Africa and countries in Latin America. He was a philanthropist of 
note, being responsible for a wing at Chicago's Mercy Hospital and the Convention 
Center and Robarts Sports Arena in Sarasota. He was interested in the arts, especially 
theater, and he and his wife coproduced a number of plays at the Asolo State Theater 
in Sarasota. But, his abiding interest was in lepidopterology, and he amassed a huge 
collection of these insects. Finally he decided that the chore was too much for one man 
part-time; thus began our association with him in 1968. 

Arthur Allyn’s interest in Lepidoptera and his desire to establish a quality institution 
for their study led to the formation of the present-day Allyn Museum of Entomology. 
During the early days of the Museum, Dr. Allyn continually purchased material that 
was needed to enhance the Museum collections, often obtaining entire collections or 
entire season's catches from people throughout the world. Later, collections or individual 





76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


specimens were donated to the museum, and less material was purchased, but the growth 
of the collection continued to its present size of about 850,000 specimens from throughout 
the world. Dr. Allyn insisted that the collection be worldwide and accepted the necessity 
of long series of specimens to show individual variation within taxa. 

In 1973 the Museum moved to its present home, and it became apparent that it would 
outgrow the new collection range rapidly if something were not done. Dr. Allyn had 
been impressed by the compactor housing the collections at the Missouri Botanical Gar- 
dens, and we immediately decided that the museum needed such housing. Thus was 
installed the first entomological compactor in the United States, a system that served as 
the model for similar systems elsewhere in this hemisphere. It was an excellent choice. 

Concern about the backlog of papers in many lepidopterological journals led to the 
founding of the Bulletin of the Allyn Museum late in 1971. This publication, patterned 
after American Museum Novitates, has gone through 105 numbers as of this writing, 
with indices after every 20 numbers. The Bulletin rapidly evolved into a refereed pub- 
lication, but final responsibility for the content of numbers rests with individual authors. 

Dr. Allyn became an accomplished scanning electron microscopist largely through his 
own efforts. One of his photographs was featured on the cover of an issue of Annals of 
the Entomological Society of America. After studying a number of entomological papers 
with which he disagreed, based on his knowledge of the physical sciences, he began to 
publish papers on structures of Lepidoptera not only with Museum staff, but also with 
such authors as Dr. John C. Downey, Dr. Miriam Rothschild and Professor David Spencer 
Smith. Other papers were in varying degrees of completion at his death, some of which 
may appear in future Bulletin numbers. Those papers that were published are standards 
for the field, and it is a tribute to Arthur Allyn and his coauthors that they are frequently 
cited not only in lepidopterological, but also in sources oriented toward scanning electron 
microscopy. Dr. Allyn also cooperated with various researchers in other fields, and his 
photographs have appeared in publications on optic, muscular and nervous systems of 
both vertebrates and invertebrates. A complete listing of Dr. Allyn’s publications is given 
in the Bulletin of the Allyn Museum, number 97. 

Eventually, Dr. Allyn began to search for an orderly transition from the independent 
nature of the Allyn Museum of Entomology to a more structured, but secure status to 
assure its permanence. After examining many options, he presented the Board of Direc- 
tors of the Allyn Museum with a proposal from the University of Florida Foundation 
which was accepted, and the Museum became a part of the Florida State Museum in 
1981. Eventually the present facility will be moved to Gainesville, and the Museum’s 
metamorphosis will be complete. 

Arthur Allyn was a benefactor of the Lepidopterists’ Society in numerous ways. In the 
late 1960's, the Society faced severe financial difficulties from which Dr. Allyn rescued 
it in return for financial accountability from the officers. Through a series of excellent 
treasurers, the Society has managed to remain a viable entity ever since. Equally impor- 
tant was the establishment of the Karl Jordan Medal for papers of lepidopterological 
excellence. There have been ten Jordan Medal awards since 1973, and the award winners 
have been truly international. Not only is the United States represented by Jordan Medal 
laureates, but also France, Canada and England. 

Those of us who were close to Arthur Allyn will miss him for his generosity, excellent 
judgment and common sense. He would not want this, however, to become an overriding 
emotion: he would demand that we continue as before. Lepidopterology, along with 
many other pursuits, is better for its association with him. 


LEE D. MILLER AND JACQUELINE Y. MILLER, Allyn Museum of Entomology of the 
Florida State Museum, 3621 Bay Shore Road, Sarasota, Florida 33580. 


Journal of the Lepidopterists’ Society 
40(1), 1986, 77-78 


LIONEL GEORGE HIGGINS (1891-1985) 


With the recent death of Lionel Higgins, at the age of 94, one of our few remaining 
links with pre-War entomology has been severed. Perhaps reflecting those more leisurely 
times, in common with many of his generation he was a ‘generalist’ of distinction, 
equally at home in art and music as in his chosen profession of medicine, or indeed 
entomology. Following rheumatic fever in childhood, Lionel was pronounced too delicate 
for a formal school education. As a result, the interests that so enlivened his lifetime were 
probably kindled. Taking a medical degree at Clare College, Cambridge, he qualified at 
St. Thomas’ Hospital before serving in the 1914-18 War as a surgeon-lieutenant. Spe- 
cializing in gynaecology and obstetrics, he practiced from 1922 onwards in Woking, 
Surrey, where he continued to live after his retirement. 

In the field of Lepidoptera, he is, of course, best known for his collaborative book with 
the late N. D. Riley, “A Field Guide to the Butterflies of Britain and Europe” (1970), a 
standard work translated into at least nine languages, with world-wide sales approaching 
200,000; and its companion volume “The Classification of European Butterflies’ (1975). 
These works were the culmination of fifty years of serious study. While this is not the 
place to include a full bibliography, C. R. Smith (Type specimens of the taxa described 
by L. G. Higgins in the British Museum (Natural History), in preparation) lists more 
than 70 titles, which truly reflects his contribution since 1924, when the first work ap- 
peared. Probably he would have regarded the Melitaeinae as “his” group. Major contri- 
butions in 1941, 1950, 1955, 1960, 1978 and 1981 provided a firm foundation for future 
work. Tangible recognition of his contributions to natural history include the Stamford 
Raffles Award (Zoological Society of London) and the H. H. Bloomer Award (Linnean 
Society). 

As well as being a prolific author, he was an indefatigable collector, both of butterflies 
and books. Accompanied by his wife, Nesta, he collected extensively in the holarctic 





78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


region. As recently as 1978, shortly before his wife’s sad and unexpected death, they 
were to be found wielding their nets in the mountains of Kashmir! The British Museum 
(Natural History) is the direct beneficiary—his incomparable collection in excess of 30,000 
specimens was generously bequeathed to this Institution. As for his library, much passed 
to the Hope Entomological Collection, Oxford; but over the years, many volumes were 
presented to the BM(NH), including several rare works by Jacob Htibner. 

Despite Lionel Higgins’ eminence, it must be acknowledged that some of his ideas 
attracted criticism. Aware of this, he maintained a dignified, but never dogmatic, con- 
fidence in his convictions. In methodology, he was a man of his times (few of us could 
claim to be anything else!). While the numerical and cladistic revolutions passed him by, 
he remained true to his basic principles—overall similarity and the equivalence of species- 
groups and genera. It is impossible to predict how the future will judge his work, but 
personal experience suggests that the results of “traditional” methods should be treated 
with the utmost respect. 

How can the loss of such a man be measured? A glance at his correspondence file 
gives some indication. There can hardly be a lepidopterist of note who has not com- 
municated with him over some problem. All were given the benefit of his lifetime’s 
experience. The personal touches perhaps give most indication of the esteem and affec- 
tion in which he was held: photographs sent by correspondents, showing themselves and 
sometimes their families, at ease in their homes. For me, the abiding memory will be of 
a battered briefcase, a cork postal-box containing the latest treasure from the Pamirs or 
Urals, and above all the half serious admonition, always delivered with a twinkle in the 
eye, that I really should learn something about palaearctic butterflies. 


LITERATURE CITED 


Hiccins, L. G. 1941. An illustrated catalogue of the Palearctic Melitaea (Lep. Rhopa- 
locera). Trans. Roy. Entomol. Soc. London 91:175-365. 

1950. A descriptive catalogue of the Palaearctic Euphydryas (Lepidoptera: 

Rhopalocera). Trans. Roy. Entomol. Soc. London 101:435-—489. 

1955. A descriptive catalogue of the genus Mellicta Billberg (Lepidoptera: 

Nymphalidae) and its species, with supplementary notes on the genera Melitaea and 

Euphydryas. Trans. Rey. Entomol. Soc. London 106:1-131. 

1960. A revision of the melitaeine genus Chlosyne and allied species (Lepi- 

doptera: Nymphalinae). Trans. Roy. Entomol. Soc. London 112:381-467. 

1975. The classification of European butterflies. Collins, London. 

1978. A revision of the genus Euphydryas Scudder (Lepidoptera: Nymphali- 

dae). Entomol. Gaz. 29:109-115. 

1981. A revision of Phyciodes Hiibner and related genera, with a review of 
the classification of the Melitaeinae. Bull. Brit. Mus. (Nat. Hist.) (Entomol.) 43:77- 
243. 

Hiccins, L. G. & N. D. Ritey. 1970. A field guide to the butterflies of Britain and 
Europe. Collins, London. 




















PHILLIP R. ACKERY, Butterfly Section, British Museum (Natural History), Cromwell 
Road, London SW7 5BD, England. 


Date of Issue (Vol. 40, No. 1): 9 October 1986 








EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
Boyce A. DRUMMOND III, DOUGLAS C. FERGUSON, THEODORE D. SARGENT 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Short manuscripts concerning new state records, current events, and notices 
should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence, KS 66044 
U.S.A. Journal contributors should prepare manuscripts according to the following in- 
structions. 

Abstract: An informative abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name with author. Mea- 
surements should be given in metric units; times should be given in terms of the 24- 
hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. References 
to footnotes should be numbered consecutively, and the footnotes typed on a separate 
sheet. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In general notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30). 

Illustrations: Only half of symmetrical objects such as adults with wings spread 
should be illustrated, unless whole illustration is crucial. Photographs and drawings should 
be mounted on stiff, white backing, arranged in the desired format, allowing (with 
particular regard to lettering) for reduction to their final width (usually 4% inches). 
Illustrations larger than 8% x 11 inches are not acceptable and should be reduced pho- 
tographically to that size or smaller. The author’s name, figure numbers as cited in the 
text, and an indication of the article’s title should be printed on the back of each mounted 
plate. Figures, both line drawings and halftones (photographs), should be numbered 
consecutively in Arabic numerals. The term “plate” should not be employed. Figure 
legends must be typewritten, double-spaced, on a separate sheet (not attached to the 
illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to 
each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the 
proofs. A page charge of $60 per full Journal page is applicable to all authors. Unfunded 
authors may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


PRESIDENTIAL ADDRESS, 1984: A TRIBUTE TO THE AMATEUR. 
Lee D. Miller) jo 
A NEw SPECIES OF EPIDROMIA (NOCTUIDAE) FROM FLORIDA. 
M. Alma Solis 2.00 i eC 


THE LARVA AND PUPA OF LYCOREA PIETERI LAMAS (DANAIDAE). 
David Kenneth Wetherbee 00 


LIFE HISTORY AND HABITS OF EXOTELEIA ANOMALA HODGES, A 
PONDEROSA PINE NEEDLE MINER IN THE SOUTHWESTERN 
UNITED STATES (GELECHIIDAE). Robert E. Stevens ................ 


BIOLOGY AND IMMATURE STAGES OF HEMILEUCA DIANA AND H., 
GROTEI (SATURNIIDAE). Paul M. Tuskes ae 


NATURAL HISTORY AND ECOLOGICAL CHEMISTRY OF THE NEO- 
TROPICAL BUTTERFLY PAPILIO ANCHISIADES (PAPILIONIDAE). 
Allen M. Young, Murray S. Blum, Henry M. Fales & Z. 
Bian oe a 


PYRGUS COMMUNIS AND P. ALBESCENS (HESPERIIDAE) IN NEVADA. 
George. T. Austin: 0 


First REPORTED MALES, SPECIES STATUS, AND AFFINITIES OF 
EPARGYREUS SPANNA EVANS (HESPERIIDAE). Kurt Johnson 
dx David Matusik 00 ee 


GENERAL NOTES 
Tryon Reakirt: A Sequel: F. Martin Brown 1... 


Saturnia walterorum (Saturniidae) in Mexico: A New National Record. Kirby 
L. Wolfe & Marvin 'D.: Valverde ue 


Emergence of Adult Ectomyelois muriscus (Dyar) (Pyralidae) from a Pod of 
Theobroma simiarum Donn. Smith (Steruliaceae) in Costa Rica. Allen 
M. Young te i OO CAA ee 


The Female of Papilio xanthopleura Godman & Salvin (Papilionidae). Kurt 
Johnson, Rick Rozycki\<> David, Matusik) 0000) 


Placement and Fate of Monarch Butterfly Pupae in Northern California. 
Leslie V. Smith, so oats 


Aberrant Postmedial Spots in Icaricia icarioides lycea and I. acmon (Lycaen- 
idae), Richard Holland i 


Notes on a Costa Rican “Monkey Slug’ (Limacodidae). Allen M. Young ... 
Book REVIEWS 
OBITUARIES 


TARR Ren nee e nee nen ene en nnn e eee naan nen n anne Rane nnn en aeenenaeanennnennanenananenaeeensanaaennnann sana aanenestenaeeeennsenaneneneeansanceneneneannaee 


ANNOUNCEMENT 
Inauguration of Manuscript Dating in the Jourrral cc ccsesssseeevecssuceeneresneeie 


Volume 40 1986 Number 2 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





23 October 1986 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


DOUGLAS C. FERGUSON, President OLAF H. H. MIELKE, Vice President 
CLIFFORD D. FERRIS, Immediate Past President EBBE SCHMIDT NIELSEN, Vice Pres- 
GERARDO LAMAS M., Vice President © ident 

RICHARD A. ARNOLD, Secretary ERIC H. METZLER, Treasurer 


Members at large: 


BoyYcE A. DRUMMOND III MIRNA M. CASAGRANDE M. DEANE BOWERS 
JOHN LANE EDWARD C. KNUDSON RICHARD L. BROWN 
ROBERT K. ROBBINS FREDERICK W. STEHR PAUL A. OPLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Affiliate members—annual dues $5.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 48229, U.S.A.; and address changes to: 
Julian P. Donahue, Assistant Secretary, Dept. of Entomology, Los Angeles County Mu- 
seum of Natural History, 900 Exposition Blvd., Los Angeles, CA 90007 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Publications Coordinator. 
The Commemorative Volume is $6; for back issues, see the NEWS for prices or inquire 
to Publications Coordinator: Ronald Leuschner, 1900 John St., Manhattan Beach, Cali- 
fornia 90266 U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, % Richard A. Arnold, Secretary, 50 Cleaveland Rd. #3, Pleasant 
Hill, CA 94523. Second-class postage paid at Los Angeles, CA and additional mailing 
offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, Julian P. 
Donahue, Assistant Secretary, Dept. of Entomology, Los Angeles County Museum of 
Natural History, 900 Exposition Blvd., Los Angeles, CA 90007 U.S.A. 


Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from 
Dyar 1899, J. New York Entomol. Soc. 7: 61-67. Suggested by Marc E. Epstein. 


JouRNAL OF 
Tue LeEpIpoPTERISTS’ SOCIETY 


Volume 40 1986 Number 2 


Journal of the Lepidopterists’ Society 
40(2), 1986, 79-92 


WHY PIERIS RAPAE IS A BETTER NAME 
THAN ARTOGEIA RAPAE (PIERIDAE) 


ROBERT K. ROBBINS 


Department of Entomology, MRC NHB 127, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560 


PAMELA M. HENSON 


Smithsonian Archives, A & I 2135, Smithsonian Institution, 
Washington, D.C. 20560 


ABSTRACT. We show that there is no phylogenetic justification for changing the 
name of Pieris rapae to Artogeia rapae. We “define” Pieris by the presence of andro- 
conial basal lobes, and suggest that this grouping, which includes P. rapae, P. brassicae, 
and P. napi, is monophyletic. Female genital characters indicate that Perrhybris, Ita- 
ballia, and Ganyra are the closest relatives of Pieris. We discuss criteria for choosing 
generic nomenclature, and suggest that the following guidelines will best promote no- 
menclatural stability. If a genus is monophyletic, do not change the name. If a genus is 
not monophyletic, choose the combination of monophyletic generic groupings that will 
create the fewest name changes. If another option causes more name changes now but 
will be more stable in the future because of better evidence for monophyly, then present 
the reasons and evidence for that choice. 


Pieris rapae Linnaeus is one of the best known and commonly en- 
countered temperate area butterflies. Although native to the Palaearc- 
tic, it is now nearly ubiquitous in suitable disturbed habitats in North 
America (Howe 1975), New Zealand (Gibbs 1980), and Australia 
(Common & Waterhouse 1981). Because P. rapae is widely distributed, 
easily reared, and a pest on cultivated crucifers, it has been extensively 
studied in the agricultural, ecological, and physiological literature 
(Harcourt 1966, Dempster 1969, Aplin et al. 1975, Slansky & Feeny 
1977, Blau et al. 1978, Kobayashi & Takano 1978, Yamamoto & Ohtani 
1979, Wolfson 1980, Chew 1981, Jones et al. 1982, Gilbert 1984, Ma- 
guire 1984). 

The generic placement of P. rapae has recently been changed from 
Pieris to Artogeia Verity. Schrank (1801) placed rapae in Pieris when 
he originally described the genus, and Klots (1933) retained this generic 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


placement in his systematic treatment of world Pieridae. Verity (1947) 
proposed Artogeia as a subgenus including rapae, and Kudrna (1974) 
and Higgins (1975) elevated it to generic rank, an action that has been 
followed in some general works (Pyle 1981, Miller & Brown 1981) but 
not others (Kawazoé & Wakabayashi 1976, Opler & Krizek 1984). This 
situation was further complicated when Kudrna later treated Artogeia 
as a subgenus (Blab & Kudrna 1982), and Feltwell and Vane-Wright 
(1982) suggested that Artogeia might not be monophyletic. 

In this paper we assess the evidence for switching rapae from Pieris 
to Artogeia. We took up this project because of repeated inquiries from 
scientists in a variety of biological disciplines to the National Museum 
of Natural History concerning the proper generic name for rapae. We 
address the following questions: ““What is the evidence for the change 
in generic nomenclature and is it compelling?’”’, ““What is the ‘best 
definition’ for Pieris?’’, “What are the closest relatives of Pieris?’’, and 
“What criteria will promote stability of generic nomenclature?” 

It is not our intent to produce a definitive work on Pieris systematics. 
Besides reporting the results of a few representative female genitalic 
dissections, we discuss published information only, all of which was 
available to Kudrna and Higgins, with the exception of two recent 
papers on isozymes. We discuss characters sequentially, note their states 
and distributions, and generally limit our discussion to those species 
for which we have information. Many species level decisions, partic- 
ularly in the P. napi group, are controversial (Warren 1961, Bowden 
1972, Eitschberger 1983, Geiger & Scholl 1985); we avoid entering the 
fray because it is largely irrelevant to our purpose. Finally, we show 
that treatment of all Pieris species would not alter our conclusions. 


PIERIS RAPAE OR ARTOGEIA RAPAE 


In this section, we ask whether rapae is more closely related to napi 
Linnaeus—the type of Artogeia—or to brassicae Linnaeus—the type 
of Pieris. The classification of Kudrna (1974) and Higgins (1975) im- 
plies that the former is correct, while others (Mariani 1937, Geiger 
1981, Geiger & Scholl 1985) suggest the opposite. These two possibil- 
ities are represented by alternative phylogenies (Figs. 1 & 2). 

To determine primitive character states among these species (the 
state at point A in the phylogenies), we used two sets of outgroup 
species. The first set is Pontia daplidice Linnaeus and Synchloe callid- 
ice Hubner. Pierid specialists (Klots 1933, Bernardi 1947) considered 
them to be closely related to the brassicae-napi complex, and some- 
times included them in Pieris. Kudrna (1974) placed them next to 
Pieris and Artogeia. They are the immediate outgroups of the brassi- 
cae-napi lineage in dendrograms constructed from isozyme data (Gei- 


VOLUME 40, NUMBER 2 81 


of of 
> & > Kj 
Yr eg ¢ Yr Oa 
$ ¥ * $ & v 
C c 
B B 
A A 


1 2 


Fics. 1 & 2. Phylogenies showing cladogenesis among Pieris brassicae, P. rapae, and 
P. napi. The letters designate ancestral species in the branching sequence. 


ger 1981, Geiger & Scholl 1985). The second outgroup set is Ganyra 
Billberg, Itaballia Kaye, and Perrhybris Hiibner (sensu Klots 1938). 
We discovered that they share female genitalic characters with the 
brassicae-napi complex (detailed below), and may be more closely 
related to them than has been previously realized. 

The first character that Kudrna (1974) and Higgins (1975) used in 
their taxonomic analysis was androconial structure (illustrations in Dix- 
ey 1910, 19382, Bernardi 1947, Warren 1961). There are four major 
shapes in the “Pieris group” with slight quantitative interspecific vari- 
ation within each type: P. brassicae has one type of androconium (Fig. 
3), rapae and A. napi a second (Figs. 4 & 5), outgroups P. daplidice, 
Perrhybris, and Itaballia a third (Figs. 6 & 7), outgroup Ganyra a 
fourth (Fig. 8), and outgroup S. callidice lacks androconia. On either 
phylogeny this distribution of character states can be explained, no 
matter which outgroup state occurred at point A, by the rapae-napi 
androconium evolving at point B and the brassicae androconium evolv- 
ing at point C. Although there are other equally parsimonious possi- 
bilities, either phylogeny could produce the distribution of character 
states simply—each androconium type evolved once. Thus, although 
rapae and A. napi share a similar androconial structure, this distribu- 
tion provides no evidence for choosing between the phylogenies in 
Figs. 1 and 2. 

Kudrna (1974) and Higgins (1975) also used male genitalic charac- 


82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


W 


4 4 
3 cig 5 ] 8 


Fics. 3-8. Outlines of androconia, redrawn from Dixey (1932). Arrows in first three 
figures point to the right basal lobe. 3, Pieris brassicae; 4, P. rapae; 5, P. napi; 6, Pontia 
daplidice; 7, Itaballia demophile; 8, Ganyra josepha. 


oo 


ters for defining Artogeia. The male genitalia of A. napi and rapae 
are similar to each other, and differ from P. brassicae (illustrated in 
Klots 1933, Bernardi 1947, Kudrna 1974, Higgins 1975). The penis of 
P. brassicae has a dorsal hump and each valva has a distal pointed 
process while rapae and A. napi lack the dorsal hump and the process. 
The outgroups, like rapae and A. napi, lack a penial dorsal hump and 
process on the valva (except for Ganyra, which has a differently shaped 
valva process). Thus, the P. brassicae penis and valva morphology is 
derived, defines only P. brassicae (evolved at point C on either phy- 
logeny), and provides no information about the phylogenetic position 
of rapae. 

Higgins (1975) also used haploid chromosome numbers to differen- 
tiate Pieris from Artogeia. Reported haploid chromosome numbers (De 


VOLUME 40, NUMBER 2 83 





Fics. 9-14. Right dorsolateral view of the corpus bursae and anterior portion of the 
ductus bursae (except for Pontia callidice, which is a dorsal view with an additional 
lateral aspect of the cervix). 9, Pieris napi; 10, P. rapae; 11, P. brassicae; 12, Perrhybris 
pyrrha; 13, Ganyra josepha; 14, Pontia callidice. 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Lesse 1967, 1970, De Lesse & Brown 1971, Robinson 1971) are P. 
brassicae—15, rapae—25-26, A. napi—25-28, and for the outgroups 
P. daplidice—26, S. callidice—26, Itaballia—25-26, and Perrhybris— 
27-29. We infer that the lower haploid chromosome number of P. 
brassicae is a derived character state that defines only P. brassicae—it 
evolved at point C in the phylogenies—and that provides no infor- 
mation on the systematic position of rapae. 

Since the characters used by Kudrna (1974) and Higgins (1975) pro- 
vide no evidence for choosing between the phylogenies in Figs. 1 and 
2, the placement of rapae in Artogeia was phylogenetically unjustified. 
We now ask whether other published characters provide information 
on the generic placement of rapae. 

Mariani (1937) and Bernardi (1947) examined “Pieris” female gen- 
italia, and reported interspecific variation in morphology of the single 
signum (often called “lamina dentata” in the pierid literature) on the 
corpus bursae. The signum of A. napi has a long posterior process 
(“flagello” of Mariani, “‘tail’’ of Chang [1963]) that is lacking in rapae 
and P. brassicae and in all outgroup species (Figs. 9-14, figures in 
Mariani and Bernardi). Thus, the posterior process is a derived char- 
acter state that apparently evolved once on the lineage leading to A. 
napi, and does not give us information with which to distinguish the 
phylogenies in Figs. 1 and 2. 

Geiger (1981) and Geiger and Scholl (1985) electrophoresed enzymes 
from species of Pieris, Artogeia, Pontia, and Synchloe, but not for the 
other outgroups. They obtained a dendrogram of relationships by using 
an unweighted pair-group average clustering method on genetic sim- 
ilarities. They found that rapae is more similar to P. brassicae than to 
A. napi and that all three are more similar to each other than to Pontia 
and Synchloe. This result supports the phylogeny depicted in Fig. 1. 
There are numerous methods for coding and analyzing electrophoretic 
data, and Mickevich and Mitter (1981) propose criteria for judging 
different methods. Before uncritically accepting their dendrograms, we 
would want to know if other methods of coding and analysis corrob- 
orate their results. 

In summary, analysis of published characters indicates that the use 
of Artogeia as a genus or subgenus including rapae is phylogenetically 
unjustified. Although the male genitalia, androconia, and haploid chro- 
mosome numbers of rapae are more similar to A. napi than to P. 
brassicae, the opposite relationship is true with regard to the female 
genitalia and isozymes. Further, these similarities are based on primi- 
tive character states, as Feltwell and Vane-Wright (1982) had predict- 
ed, and do not provide the information necessary to choose between 
the phylogenies in Figs. 1 and 2. Characters of “Pieris” immature 


VOLUME 40, NUMBER 2 85 


stages may provide the information necessary to decide this point, but 
have not been used in Pieris revisions. 


THE GENUS PIERIS 


Since rapae is such a widely known species and since Kudrna and 
Higgins’ concept of Pieris and Artogeia leave rapae without certain 
generic placement, we ask in this section whether there are other, more 
reasonable definitions for Pieris. Klots (1933) revised the world pierid 
fauna. Although he narrowed the definition of Pieris—it previously 
had been a catchall genus for many questionably related pierines— 
subsequent authors have split the genus further. We ask whether any 
of these groupings are monophyletic. For outgroup comparisons, we 
use those genera that Klots considered to be most closely related to 
Pieris: Leptophobia Butler, Itaballia, Perrhybris, Ascia Scopoli (in- 
cluding subgenus Ganyra), Tatochila Butler, Phulia Herrich-Schaffer, 
and Baltia Moore. 

Three different concepts of Pieris besides that of Kudrna and Hig- 
gins have been used since 1933. For ease of communication, we list 
representative species for each grouping, and refer the reader to the 
original work for a complete list. Klots (1933) placed brassicae, rapae, 
napi, callidice, daplidice, and pylotis in Pieris. Mariani (1937) and 
Bernardi (1947) put the first four of these species in Pieris, while Hig- 
gins and Riley (1970) restricted Pieris to the first three. (Note that 
Higgins [1975] later narrowed the genus further, like Kudrna, to in- 
clude only brassicae and close relatives.) 

Klots (1933) defined Pieris with a paragraph of character states. For 
the most part, however, they are too ambiguous to code accurately. 
For example, how does one code “antenna long, with abrupt club” 
(Pieris), “antenna long, with usually somewhat abrupt club” (Ascia), 
and “antenna long, with flattened abrupt club” (Tatochila)? Further, 
each of Klots’ generic character states is shared with at least one out- 
group genus. Because of character state ambiguity and the lack of 
unique, potentially defining character states, we found no evidence in 
Klots’ work to indicate that his concept of Pieris is monophyletic. 

Mariani (1937) and Bernardi (1947) apparently ignored pylotis (a 
neotropical species that does not “look” like other Pieris species) and 
moved daplidice to Pontia. Pontia daplidice has forewing veins R; and 
R,,; fused while they are separate in the other Pieris species. Outgroups 
Phulia and Perrhybris have the fused veins while the other outgroups 
have separate veins (Klots 1933). Because both character states are 
found in the outgroups, the primitive character state is ambiguous. 
Phylogenetic interpretation of this character is thus equivocal. 

Higgins and Riley (1970) joined callidice with daplidice in Pontia, 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


so that their Pieris grouping consisted of brassicae, rapae, napi, and 
close relatives. The androconial lateral edges of these species and their 
close relatives protrude basally to form lobes (Dixey 1932, Bernardi 
1947, Warren 1961) (Figs. 83-5) while the androconial basal edge is flat 
in daplidice and relatives (Bernardi 1947) (Fig. 6), pylotis (Dixey 1932), 
and all outgroup genera (Dixey 1932) (Figs. 7 & 8). Since daplidice 
and pylotis share the primitive state—that which occurs in the out- 
group genera—the androconial basal lobes would appear to be a de- 
rived, defining character for the brassicae-napi complex. This situation 
contrasts with the one in the previous section, in which rapae and napi 
share an androconium type that does not reflect phylogenetic relat- 
edness because brassicae does not possess the primitive character state. 

We “define” Pieris by the androconial basal lobes. Although we are 
reluctant to define a genus on the basis of one character state, there 
are no alternatives in this case. From published illustrations of andro- 
conia (Dixey 1932, Bernardi, 1947), we place the following specific 
taxa—listed in Bernardi—in Pieris: brassicae, deota de Niceville, bras- 
sicoides Guerin, krueperi Staudinger, tadjika Groum-Grshimailo, ca- 
nidia Sparrman, manni Mayer, rapae, dubernardi Oberthur, extensa 
Poujade, stoetzneri Draeseke, napi, virginiensis Edwards, ochsenhei- 
meri Staudinger, ergane Geyer, melete, and davidis Oberthur. Al- 
though there are other taxa, particularly in the napi group, that are 
given specific rank by some authors, we leave species level decisions 
to others. 

We believe that this grouping is the most reasonable and stable one 
for Pieris. There is an enormous biological literature on Pieris brassi- 
cae, Pieris rapae, and Pieris napi, and the name Pieris is widely rec- 
ognized by nontaxonomists in connection with these species. Our 
grouping will preserve this association, and because it is based on the 
best available evidence for monophyly, it is most likely to be stable in 
the future. 

There are three morphologically distinct groups within Pieris. The 
P. brassicae group (brassicae, deota, brassicoides) has the androconial 
and male genital structures of brassicae, and is probably a monophy- 
letic lineage defined by these structures. The P. napi group (napi, 
virginiensis, ochsenheimeri, ergane, melete, davidis, and presumably 
stoetzneri, extensa, and dubernardi—Bernardi [1947]) has a posterior 
process on the signum, which probably defines this group as a mono- 
phyletic lineage. The P. rapae group (krueperi, tadjika, canidia, man- 
ni, rapae) lacks derived character states. There is no evidence to in- 
dicate whether it is monophyletic or whether it is phylogenetically 
more closely related to the P. brassicae or P. napi groups. Thus, even 
if we had examined all Pieris species in the previous section, it would 


VOLUME 40, NUMBER 2 87 


not have provided us with evidence on the phylogenetic position of P. 
rapae. Interestingly, the same three groups result when isozyme data 
are analyzed phenetically (Geiger & Scholl 1985). 


THE RELATIVES OF PIERIS 


In this section we ask which genus or genera are most closely related 
to Pieris. From the work of Mariani (1937) and Bernardi (1947), it 
appeared that the bursa copulatrix, particularly signum location and 
shape, had states that might provide information on the phylogenetic 
position of Pieris. Because this character was promising, but unrecord- 
ed for many of the outgroups, we dissected the female genitalia of 
species in Pieris and related genera. 

We recorded three character states of the bursa copulatrix. In the 
first, the signum is a narrow transverse band located at the posterior 
end of the corpus bursae just around the entrance to the ductus bursae 
(Fig. 14). We recorded this state in Pontia (daplidice, protodice), 
Synchloe (callidice), Leptophobia (eleone Hewitson, aripa Boisduval), 
and Ascia (monuste Linnaeus). It also occurs in Tatochila, Phulia, 
Baltia, and close relatives (Field 1958, Herrera & Field 1959, Field & 
Herrera 1977), in the pierine Aporia Htibner (Mariani 1937) and the 
coliadines Colias Fabricius (Mariani 1937) and Eurema Hubner (Field 
1950). 

In the second character state, the signum is located on the right 
dorsolateral side of the corpus bursae well anterior to the entrance of 
the ductus bursae (Figs. 9-13). Signum shape varies, particularly in 
how far it extends posteriorly and in the amount of sclerotization of 
the median line. We recorded this character state in Pieris (brassicae, 
rapae, napi, melete), Ganyra (josepha Godman & Salvin, limona 
Schaus), Itaballia (demophile Linnaeus, viardi Boisduval, pisonis Hew- 
itson), and Perrhybris (pyrrha Fabricius, pamela Cramer [=lypera Kol- 
lar], lorena Hewitson). Mariani (1937) noted its occurrence in all 12 
Pieris species that he examined. 

A third character state is limited to Glennia pylotis. There is no 
signum. The corpus bursae and ductus bursae are greatly modified into 
a long tube that occupies the length of the abdomen. This tube grad- 
ually increases in diameter anteriorly, and the usual abrupt change in 
size that distinguishes the corpus from the ductus is absent. 

The closest relatives of Pieris appear to be Perrhybris, Itaballia, and 
Ganyra. The position of the signum on the right dorsolateral side of 
the corpus bursae is an unusual character state that is apparently re- 
stricted to these four genera. The other genera that Klots (1933) placed 
near Pieris have the signum at the posterior end of the corpus bursae, 
which is probably the primitive state for the pierines because it is also 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


found in the coliadines Eurema and Colias. A definitive survey of the 
distributions of female genital structures is obviously desirable. 

Those holarctic species that Klots (19383) put in Pieris, but which 
have recently been placed in Pontia (Higgins 1975, Miller & Brown 
1981) are often considered to be close relatives of Pieris. However, we 
know of no evidence that this group is more closely related to Pieris 
than to other genera, such as Tatochila, Phulia, and relatives (Shapiro 
1979). Further, we have found no. published characters to determine 
whether this group is monophyletic. In short, there is a glaring need 
for a worldwide treatment of the pierines. 

The placement of Glennia pylotis remains a problem. Although 
Klots (1933) treated Glennia as a subgenus of Pieris, it lacks the an- 
droconial basal lobes and signum of Pieris. There is currently no evi- 
dence to decide whether the divergent female genitalia of Glennia 
evolved from the Pieris type or from the Pontia type. 


STABILITY AND GENERIC NOMENCLATURE 


In this section we use the confusion over the generic nomenclature 
of P. rapae as an example to discuss the relationship between taxonom- 
ic method and nomenclatural stability. 

We suggest that butterfly generic nomenclature can be more objec- 
tively chosen than in the past by using the criteria of “stability” and 
“monophyly”. The Preamble to the International Code of Zoological 
Nomenclature (Int. Comm. on Zool. Nomenclature 1985) states that 
“.. the object of the Code is to promote stability and universality in 
the scientific names of animals ....’’ Ehrlich and Murphy (1982) dis- 
cuss the widespread support for a stable generic nomenclature. 

By monophyly, we refer to taxa defined by derived characters. As 
Jordan (1898) noted, “... we have here an instructive illustration of 
the fact—so very often entirely disregarded in classificatory work— 
that the presence of the same character in two different [taxa] .. . is, 
evidence of closer relationship only, if the character is a specialisation 
and not of the ancestral type.”’ Jordan’s logic is simple, but has been 
largely ignored by butterfly systematists. 

The application of stability and monophyly to groups with an estab- 
lished generic nomenclature, such as the bulk of the butterflies, is 
straightforward. If a genus is monophyletic, do not change the name. 
If a genus is not monophyletic, choose the combination of monophy- 
letic generic groupings that will create the fewest name changes. If 
another option causes more name changes now but will be more stable 
in the future because of better evidence for monophyly, then present 
the reasons and evidence for that choice. 

Ehrlich and Murphy (1982) suggested that the concept of balance 


VOLUME 40, NUMBER 2 89 


(the equivalence of categorical rank in related taxa, sensu Mayr 1969) 
also be used to decide generic nomenclature. Despite Mayr’s discussion 
of how balance might be applied, this method is subjective, particularly 
since it is unclear exactly what the method is supposed to estimate. 
Although objectivity is not itself justification for using a criterion, we 
believe that an obviously subjective one, such as balance, will promote 
instability of butterfly generic nomenclature. 

Kudrna (1974) and Higgins (1975) used the criterion of “‘similarities 
and differences” to justify their recognition of Artogeia, and did not 
mention stability and monophyly. For example, Higgins (1975) stated, 
“Their [Artogeia] genitalia, androconial scales and chromosome num- 
bers differ from those of P. brassicae and it is not satisfactory to include 
them in the same genus.” Neither worker suggested that Pieris, as used 
by Klots (1938) or Bernardi (1947), was polyphyletic. Neither discussed 
the possible confusion that would result from changing the generic 
nomenclature of P. rapae and P. napi. 

There are many problems with the criterion of similarities and dif- 
ferences. (1) The similarities and differences used by Kudrna and Hig- 
gins do not provide information on the phylogenetic position of P. 
rapae. This example is a clear illustration that similarities and differ- 
ences alone are insufficient to establish monophyly. (2) If Kudrna and 
Higgins had examined female genitalia and isozymes (as opposed to 
male genitalia, androconia, and chromosome numbers), they would 
have put rapae in Pieris. When taxonomic conclusions depend upon 
the character set used, the result is instability. (8) We are certain that 
Kudrna and Higgins believed that Artogeia should be split from Pieris 
because it is “sufficiently different.’’ However, if one “‘authority”’ states 
that a difference is sufficient to split a genus, but another disagrees, 
then how can these conflicting views be resolved? It is evident that the 
criterion of similarities and differences promotes instability, and should 
not be used. 

Kudrna (1974) and Higgins (1975) assumed that the divergent mor- 
phology of P. brassicae is the result of phylogenetic distance, but did 
not consider that it might be the result of rapid evolution. We hypoth- 
esize that rearrangement of genes caused by extensive chromosomal 
fusion—haploid chromosome number decreased from about 26 to 15 
at point C in Figs. 1 and 2 (discussion in White 1973)—affected gene 
expression during development (the “position effect”; Dobzhansky 1957, 
White 1973), and is causally related to the divergent male genital, 
androconial, and larval (D. Weisman, pers. comm.) morphology of P. 
brassicae. Chromosomal rearrangements would not be expected to af- 
fect the protein products of structural genes, however—an expectation 
consistent with isozyme data (Geiger 1981, Geiger & Scholl 1985). Our 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


hypothesis generates the testable prediction that morphology and chro- 
mosome numbers are perfectly correlated; all species in the P. brassicae 
group should have reduced haploid chromosome numbers (about 15) 
while none in the P. rapae and P. napi groups should have the reduced 
numbers. 

J. H. Comstock (1893) wrote: “Here I believe lies the work of the 
systematist of the future. The description of a new species, genus, 
family or order, will be considered incomplete until its phylogeny has 
been determined so far as is possible with the data at hand.’’ Com- 
stock’s vision of the holarctic butterfly “systematist of the future’ is, 
by and large, still just a vision. Until we have reasonable phylogenies, 
generic nomenclature is bound to be unstable. In the meantime, sug- 
gested changes in generic nomenclature will hopefully be based on 
evidence of monophyly, and proposed with due regard for stability. 


ACKNOWLEDGMENTS 


We conducted this project under the auspices of the Maryland Center for Systematic 
Entomology, a consortium of the Department of Entomology, Smithsonian Institution, 
Department of Entomology, University of Maryland, and the Systematic Entomology 
Laboratory (USDA). We thank C. W. Mitter of the Center for his support. We thank L. 
D. Miller of the Allyn Museum of Entomology for the loan of a specimen. For providing 
thoughtful criticisms of various manuscript drafts, we thank P. R. Ackery, A. Aiello, S. 
R. Bowden, M. D. Bowers, J. M. Burns, C. J. Callaghan, F. S. Chew, S. P. Courtney, C. 
V. Covell, R. de Jong, J. C. Downey, U. Eitschberger, J. N. Eliot, L. F. Gall, J. S. 
Glassberg, R. W. Hodges, G. Lamas, C. W. Mitter, S. S. Nicolay, P. A. Opler, R. W. 
Poole, J. E. Rawlins, R. L. Rutowski, A. M. Shapiro, G. B. Small, R. I. Vane-Wright, and 
B. A. Venables. We thank G. Venable for professional illustrative aid. 


LITERATURE CITED 


APLIN, R. T., W. R. D’ARCY & M. ROTHSCHILD. 1975. Examination of the large white 
and small white butterflies (Pieris spp.) for the presence of mustard oils and mustard 
oil glycosides. J. Entomol. (A) 50:73-78. 

BERNARDI, G. 1947. Revision de la classification des espéces holarctiques des genres 
Pieris Schr. et Pontia Fabr. (Lep., Pieridae). Misc. Entomol. 44:65-80. 

BAB, J. & O. KUDRNA. 1982. Hilfsprogram fiir Schmetterlinge. Naturschutz aktuell 6: 
1-135. 

BLAU, P. A., P. FEENY & L. CONTARDO. 1978. Allylglucosinolates and herbivorous 
caterpillars: A contrast in toxicity and tolerance. Science 200:1296-1298. 

BOWDEN, S. R. 1972. Pieris napi in America: Genetic imbalance in hybrids. Proc. Brit. 
Entomol. Nat. Hist. Soc. 4:1038-117. 

CHANG, V. C. S. 1963. Quantitative analysis of certain wing and genitalia characters 
of Pieris in western North America. J. Res. Lepid. 2:97-125. 

CHEW, F. S. 1981. Coexistence and local extinction in two pierid butterflies. Am. Nat. 
18:655-672. 

Common, I. F. B. & D. F. WATERHOUSE. 1981. Butterflies of Australia. Rev. Ed. 
Australian Natural Science Library. Angus & Robertson, London. 682 pp. 

Comstock, J. H. 1893. Evolution and taxonomy; an essay on the application of the 
theory of natural selection in the classification of animals and plants, illustrated by 


VOLUME 40, NUMBER 2 91 


a study of the evolution of the wings of insects and by a contribution to the classification 
of the Lepidoptera. In Wilder quarter century book, pp. 37-113. Comstock Publish- 
ing Company, Ithaca, New York. 493 pp. 

DE LEssE, H. 1967. Les Nombres de chromosomes chez les Lépidoptéres Rhopalocéres 
Néotropicaux. Ann. Soc. Entomol. France (N.S.) 3:67-136. 

1970. Les Nombres de chromosomes chez les Lépidoptéres Rhopalocéres en 
Amérique centrale et Colombie. Ann. Soc. Entomol. France (N.S.) 6:347-358. 

DE LeEssE, H. & K. S. BROWN. 1971. Formules chromosomiques de Lépidoptéres Rho- 
palocéres du Brésil. Bull. Soc. Entomol. France 76:131-1387. 

DEMPSTER, J. P. 1969. Some effects of weed control on the numbers of the small 
cabbage white (Pieris rapae) on brussel sprouts. J. Appl. Ecol. 6:339-345. 

DixEy, F. A. 1910. The plume scales of the Pierinae. Proc. Entomol. Soc. London 1909: 
xciii—cxli. 

1932. The plume-scales of the Pierinae. Trans. Entomol. Soc. London 80: 
57-75. 

DOBZHANSKY, T. 1957. Genetics and the origin of species. 3rd ed. Columbia University 
Press, New York. 364 pp. 

EHRLICH, P. R. & D. D. MurpHy. 1982. Butterfly nomenclature: A critique. J. Res. 
Lepidop. 20:1-11. 

EITSCHBERGER, U. 1983. Systematische Untersuchungen am Pieris napi-bryoniae- 
Komplex (s.1.). Herbipoliana 1(1983)(1):I-XXII, 1-504; (2):1-601. 

FELTWELL, J. & R. I. VANE-WRIGHT. 1982. Classification of Pieris brassicae. In J. 
Feltwell, Large white butterfly. The biology, biochemistry and physiology of Pieris 
brassicae (Linnaeus), pp. 1-11. Dr. W. Junk, The Hague. 535 pp. 

FIELD, W. D. 1950. A revision of Eurema Hubner subgenus Tericolias Rober. De Acta 
Zool. Lilloana 9:359-374. 

1958. A redefinition of the butterfly genera Tatochila, Phulia, Piercolias, and 
Baltia, with descriptions of related genera and subgenera. Proc. U.S. Nat. Mus. 108 
(No. 3396):103-131. 

FIELD, W. D. & J. HERRERA. 1977. The pierid butterflies of the genera Hyposchila 
Ureta, Phulia Herrich-Schaffer, Infraphulia Field, Periphulia Field, and Piercolias 
Staudinger. Smiths. Contr. Zool. No. 232. 64 pp. 

GEIGER, H. J. 1981. Enzyme electrophoretic studies on the genetic relationships of 
pierid butterflies (Lepidoptera: Pieridae) I. European taxa. J. Res. Lepidop. 19:181- 
195. 

GEIGER, H. J. & A. SCHOLL. 1985. Systematics and evolution of holarctic Pierinae 
(Lepidoptera). An enzyme electrophoretic approach. Experientia 41:24—29. 

Gipss, G. W. 1980. New Zealand butterflies, identification and natural history. Collins, 
Auckland. 207 pp. 

GILBERT, N. 1984. Control of fecundity in Pieris rapae. III. Synthesis. J. Anim. Ecol. 
53:599-609. 

HARCOURT, D. G. 1966. Major factors in survival of the immature stages of Pieris 
rapae. Can. Entomol. 98:653-662. 

HERRERA, J. & W. D. FIELD. 1959. A revision of the butterfly genera Theochila and 
Tatochila (Lepidoptera: Pieridae). Proc. U.S. Nat. Mus. 108(No. 3408):467-514. 
Hiccins, L. G. 1975. The classification of European butterflies. Collins, London. 320 pp. 
HiccIns, L. G. & N. D. RiLEy. 1970. A field guide to the butterflies of Britain and 

Europe. The International Series. Houghton Mifflin, Boston. 380 pp. 

Howe, W. H. 1975. The butterflies of North America. Doubleday, Garden City. 633 pp. 

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1985. International 
Code of Zoological Nomenclature. U. of California Press, Berkeley. 338 pp. 

JonEs, R. E., J. R. HART & G. D. BULL. 1982. Temperature, size, and egg production 
in the cabbage butterfly Pieris rapae. Austral. J. Zool. 30:223-232. 

JORDAN, K. 1898. Contributions to the morphology of Lepidoptera. Novit. Zool. 5:374- 
415. 

KAWAZOE, A. & M. WAKABAYASHI. 1976. Coloured illustrations of the butterflies of 
Japan (in Japanese). Hoikusha Publishing Co., Osaka. 422 pp. 











92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Kiots, A. B. 1933. A generic revision of the Pieridae (Lepidoptera). Entomol. Am. 12 
(new series): 189-242. 

KOBAYASHI, S. & H. TAKANO. 1978. Survival rate and dispersal of adults of Pieris rapae 
crucivora. Jap. J. Appl. Entomol. Zool. 22:250-254. 

KupRNA, O. 1974. Artogeia Verity, 1947, Gen. Rev. for Papilio napi Linnaeus (Lep., 
Pieridae). Entomol. Gaz. 25:9-12. 

Macuire, L. A. 1984. Influence of surrounding plants on densities of Pieris rapae (L.) 

- eggs and larvae (Lepidoptera: Pieridae) on collards. Environ. Entomol. 13:464—468. 

MaRIANI, M. 1937. Anatomia e fisiologia degli organi genitali femminili delle Pieris 
Schrk. (Lepidoptera, Pieridae). Festschrift 60 Geburtstag E. Strand. Vol. III, Riga 
Universitate 3, 434-450. 

Mayr, E. 1969. Principles of systematic zoology. McGraw-Hill, New York. 428 pp. 

MICKEVICH, M. F. & C. MITTER. 1981. Treating polymorphic characters in systematics: 
a phylogenetic treatment of electrophoretic data. In V. A. Funk & D. R. Brooks 
(Eds.), Advances in cladistics. Vol 1:45-58. New York Botanical Garden, New York. 
250 pp. 

MILLER, L. D. & F. M. BRowN. 1981. A catalogue/checklist of the butterflies of 
America north of Mexico. Lepid. Soc. Mem. No. 2. 280 pp. 

OPpLeErR, P. A. & G. O. KrizEK. 1984. Butterflies east of the Great Plains. An illustrated 
natural history. The Johns Hopkins University Press, Baltimore. 294 pp. 

PYLE, R. M. 1981. The Audubon Society field guide to North American butterflies. 
Alfred A. Knopf, New York. 924 pp. 

ROBINSON, R. 1971. Lepidoptera genetics. Pergamon, Oxford. 687 pp. 

SCHRANK, F. P. 1801. Fauna boica. Durchgedachte Geschichte der in Baiern Einhei- 
mischen und Zahmen Thiere. Vol. 2, Part 1. Krull, Ingolstadt. 374 pp. 

SHAPIRO, A. M. 1979. The life histories of the autodice and sterodice species-groups of 
Tatochila (Lepidoptera: Pieridae). J. New York Entomol. Soc. 87:236—255. 

SLANSKY, F. & P. FEENY. 1977. Stabilization of the rate of nitrogen accumulation by 
larvae of the cabbage butterfly on wild and cultivated food plants. Ecol. Monogr. 
47:209-228. 

VERITY, R. 1947. Le farfaile diurne d'Italia. Vol. III. Casa Editrice Marzocco, Firenze. 
318 pp. 

WARREN, B. C. S. 1961. The androconial scales and their bearing on the question of 
speciation in the genus Pieris (Lepidoptera). Entomol. Tidskr. 82:121-148. 

White, M. J. D. 1973. Animal cytology and evolution. 8rd ed. Cambridge University 
Press, Cambridge. 961 pp. 

WOLFSON, J. L. 1980. Oviposition response of Pieris rapae to environmentally induced 
variation in Brassica nigra. Entomol. Exp. Appl. 27:223-232. 

YAMAMOTO, M. & T. OHTANI. 1979. Number of eggs laid by Pieris rapae crucivora, 
compared with P. napi nesis, in Sapporo. Kontyu 47:530-539. 


‘Journal of the Lepidopterists’ Society 
40(2), 1986, 938-96 


RESTINGA BUTTERFLIES: BIOLOGY OF 
SYNARGIS BRENNUS (STICHEL) (RIODINIDAE) 


CurRTIs J. CALLAGHAN 


Rua yeddo Fiuza 595, Petropolis, Rio de Janeiro, Brazil, and 
“Pesquisador Associado” of the Museu Nacional, Rio de Janeiro 


ABSTRACT. Synargis brennus (Stichel) isa myrmecophilous riodinine butterfly that 
inhabits the restinga, a low forest on the Brazilian coast. The larval food plant Dalbergia 
ecastophylla has nectaries attractive to ants of the genera Camponotus and Azteca. Ants 
attend the larvae, drumming on them to stimulate a secretion which they subsequently 
eat from two glands located on the eighth abdominal segment. 


This paper continues the series of studies on restinga butterflies start- 
ed with the biology of Menander felsina (Callaghan 1977). The ur- 
gency of the study of this habitat is underlined by the fact that the site 
of the latter study near Rio de Janeiro has been destroyed by a housing 
development. 

Synargis brennus (Stichel) (Fig. 1), a myrmecophilous riodinine but- 
terfly, is not an endemic restinga species, but is found mostly in this 
habitat, and forms an important element of its fauna. The butterfly 
ranges from the coast in southeast Brazil across the Planalto to the 
Amazon basin, where it intergrades with S. calyce (Felder). The habitat 
where the observations were made is called “restinga’’, and consists of 
low, scrubby, dense, woody vegetation growing along the coast. The 
vegetation and physical characteristics are summarized in Callaghan 
(1977). The observations in the present study were made over three 
months during visits to Buzios, a very dry section of coastline 170 km 
E of Rio de Janeiro (Fig. 2). 

Observations were made in the field and in the laboratory. The 
letters T and A followed by a number refer to thoracic and abdominal 
segments, respectively. 


DESCRIPTION OF IMMATURE STAGES 


Egg (Fig. 3). Rounded laterally, flattened dorsally and ventrally, giving the appearance 
of a fat tire. Color shiny bronze, with a network of small ribs forming a hexagonal pattern 
smaller around the micropyle and ventrally; top of ribs irregular, with a small tubercle 
at each intersection. Micropyle circular, depressed, with numerous small openings. Du- 
ration 9 days (N = 5). 

First instar. Length 1.8 mm, head width 0.2 mm. Form rounded dorsally, flat ven- 
trally. Head black, numerous small setae on face. Thorax light brown with black dots, 
pair of reddish, broken lines dorsally; T1 bilobed, with eight long cilia pointed cephalad, 
a spiracle on each side. T2 and T3 with three long setae laterally at base of tergum. 
Abdomen with pair of reddish lines dorsally. Four lateral setae on each segment except 
last, on which there are six long setae pointed caudad. Spiracles on segments Al—A8, 
indistinct; those on Al located ventrally, those on A2—A8 laterally. Duration 6 days 
(N = 5). 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-9. 1, Female Synargis brennus resting between ovipositions,; 2, Study site 
in restinga woods, Buzios, Rio de Janeiro, Brazil; 3, Egg shells on stem of foodplant; 4, 
Second instar attended by a Camponotus ant; 5, Ants tending third instars on host plant; 
6, Fourth instars; 7, Ant tending fifth instar; 8, Prepupa; 9, Pupa. 


Second instar (Fig. 4). Length 4 mm, head width 0.4 mm. T1 light brown with two 
black horns pointed cephalad, bearing two setae extending from the ends, and many 
small teeth on the surface; between the horns cephalad are two vibrating papillae; below 
each horn is a lateral scolus with two long setae. T2 and T3 with two maior lateral setae 
each. Al-A9 with four lateral setae on edge of tergum; A9 and A10 with tail plate and 
six long setae pointed caudad. AJl setae with numerous small spines. A8 with Newcomer's 
organs well developed as two raised slits. Duration 5 days (N = 9). 

Third instar (Fig. 5). Length 6 mm, head width 0.9 mm. Head black, thorax and 
abdomen light green dorsally, light yellow ventrally. Horns of prothorax black; anal plate 
light brown, tegument covered with small white points. Spiracles light brown. Duration 
6 days (N = 7). 


VOLUME 40, NUMBER 2 95 


Fourth instar (Fig. 6). Length 10 mm, head width 1.2 mm. Like third instar, except 
area between horns on prothorax and cephalad of anal plate dark brown. Duration 5 
days (N = 4). 

Fifth instar (Fig. 7). Length 15-19 mm, head width 1.7 mm. Like fourth instar except 
T2 and T3 with V-shaped figure dorsally pointed caudad; A8 with light brown saddle 
between the Newcomer’s organs. Duration 7 days (N = 8). 

Prepupa (Fig. 8). Length 19 mm. Like fifth instar except integument mottled brown- 
gray, white spiracles. Duration 3 days (N = 2). 

Pupa (Fig. 9). Length 15 mm, width at widest part 5 mm. Two rounded horns cephalad 
with T-shaped black spot behind them; thoracic segments with dorsal hump; abdominal 
segments wide, flat dorsally, first two widest, terminating laterally in a scolus with spiracle 
surrounded by spoon-shaped scales, two rows of similar scoli located dorsally, two to each 
segment. Pupa secured by cremaster and silk girdles. Color mottled gray-brown to green- 
ish, varying in pattern between individuals. Duration 11 days (N = 2). 


Preserved material is in the author’s collection. Larvae of S. brennus 
are similar to those of Juditha molpe. The latter differs in being lighter 
green with yellow dots dorsally, and in the smooth face and horns. 
Otherwise, the larvae are morphologically very close. The ventrally 
positioned spiracle on Al is the same in both. This suggests that S. 
brennus and J. molpe may be congeneric. 


BIOLOGY 


The foodplant of Synargis brennus is Dalbergia ecastophylla (Linn. 
& Talb) which is common throughout the restinga. The leaves are 
simple, ovate, and alternate along a woody stem, with a nectary at the 
base of each petiole. The plant grows as a vine, winding its way through 
the branches of other trees and shrubs, making the restinga all but 
impenetrable. 

Females oviposit during the afternoon on all parts of the foodplant: 
leaves, leaflets, petiole, stem at base of petiole, nectaries. They first feel 
the surface with the tip of the abdomen. The egg is laid quickly, in 
less than one minute (N = 5), the female then flying to another nearby 
leaf where she rests for a few minutes before returning to another part 
of the foodplant to oviposit again. The eggs were placed near Cam- 
ponotus ants, which surrounded the female but were not hostile. 

Newly hatched larvae fed on new leaf buds or at the nectaries, and 
these preferences were maintained throughout larval development. As 
the nectaries dried up, and the leaves became tough and leathery, the 
larvae aggregated on new plant growth. Feeding took place mainly at 
night, the larvae remaining motionless on leaves or stem during the 
day where cryptic coloration made them difficult to locate. They spent 
their time exclusively on the foodplant, resting or moving about, weav- 
ing the head from side to side as they laid down silk by which they 
secured their grip. When disturbed, they raised the front half of the 
body, flopping it about. 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


From the second instar on, larvae were attended by ants (Figs. 4, 5, 
7), either Camponotus crassus ssp., which also attends M. felsina lar- 
vae in the restinga (Callaghan 1977), or Azteca sp. I never found the 
two species of ants together on the same plant. The ants drum their 
antennae on the larval head and thorax, thereby stimulating secretion 
of honeydew by Newcomer’s organs. The ants then consumed the se- 
cretion. As with M. felsina and J. molpe, Camponotus ants protected 
the larvae by spraying formic acid at intruders (Callaghan 1982). The 
larvae possess vibratory papillae; these vibrate rapidly when they walk, 
and may attract ants. 

Larvae with no or few ants were sometimes parasitized. I discovered 
one solitary, unattended larva parasitized by an ichneumon wasp. The 
larvae are not otherwise dependent on ants. In the laboratory, I raised 
larvae from first instar to pupation without ant presence. 

Like J. molpe, S. brennus larvae are cannibalistic. On two occasions, 
when fresh food was lacking in the laboratory, the larger brennus larvae 
killed and ate smaller ones. This behavior in nature increases the chances 
of later instar larvae reaching pupation, should food resources fail. 

At the end of the fifth instar, larvae turn a mottled brown-gray, 
cease feeding, and remain motionless on the foodplant. Pupation takes 
place on the ventral surface of the leaves, and the ants lose interest. 
The imago emerges 11 days later. 


ACKNOWLEDGMENTS 


I thank Keith Brown and Woodruff Benson for their helpful comments on the manu- 
script, and Dr. Tomashiru of the Universidade Estadual de Campinas for determining 
the foodplant. 


LITERATURE CITED 


CALLAGHAN, C. J. 1977. Studies on restinga butterflies I. Life cycle and immature 
biology of Menander felsina (Riodinidae) a myrmecophilous metalmark. J. Lepid. 
Soc. 31:173-182. 

1981(82). Notes on the immature biology of two myrmecophilous Lycaenidae: 

Juditha molpe (Riodininae) and Panthiades bitias (Lycaeninae). J. Res. Lepid. 20: 

36-42. 





Journal of the Lepidopterists’ Society 
40(2), 1986, 97-106 


MALE AND FEMALE GENITALIA OF PHOEBIS EDITHA 
(BUTLER): HOW THEY DIFFER FROM HISPANIOLAN 
P. SENNAE (LINNAEUS) (PIERIDAE) 


JOHN G. COUTSIS 
4 Glykonos Street, Athens 10675, Greece 


ABSTRACT. Male and female genitalia of the Hispaniolan endemic Phoebis editha 
are figured, described, and compared with those of superficially similar P. sennae from 
Hispaniola. Results are based on 14 male and 2 female P. editha, and 17 male and 2 
female P. sennae. Males of P. editha differed from those of P. sennae in at least six 
ways, including narrower sacculus, and longer ampullary process. Females of P. editha 
differed from those of P. sennae in at least five ways, including more heavily sclerotized 
apophyses anteriores, and shorter, wider 8th tergum. These differences, together with 
the facts of sympatry, synchronism, and different larval foodplants, suggest that the taxa 
are specifically distinct and not forms of the same species. 


Phoebis editha, endemic to Hispaniola, was originally described as 
a distinct species (Butler 1870). Due to superficial similarity between 
the males of P. editha and P. sennae, the taxonomic status of the 
former has been in doubt. Most recently, D’Abrera (1981) suggested 
P. editha may represent a rare form of P. sennae. The female of P. 
editha at times has been considered a dry season form of P. sennae, 
or even of P. philea (Johansson). This latter view is reported, but not 
endorsed, by Riley (1975). 

Recently, I stated reasons why P. editha should be considered spe- 
cifically distinct from P. sennae, the most important of which were 
sympatry, synchronism, different larval foodplants, and different male 
genitalia (Coutsis 1983). Due to unavailability of material at the time, 
I was unable to illustrate the genital differences. 

It is now possible for me to describe and figure male and female 
genitalia of P. editha because I have been able to borrow two male 
and two female specimens. For comparison, genitalia of two male and 
two female Hispaniolan P. sennae are also figured. The findings agree 
with those derived from a study of 12 male P. editha and 15 male P. 
sennae, which I carried out between 1952 and 1958 while doing field 
work in Hispaniola. 

The drawings were done using a Wild M5 stereomicroscope with 
drawing tube. The appendages were studied and drawn while they 
were immersed in 80% ethyl alcohol, free from pressure due to slide 
mounting, and thus free from distortion. 

The genital terminology used is based on Tuxen (1970) and Higgins 
(1975). 


98 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Dorsal Margin 
Dorsal Process 


Distal Margin 










<., Proximal Margin 


ee S fie Ampullary Process 





Apex i 
Sacculus 








1339 








Fic. 1. Male Phoebis editha from Port-au-Prince, Haiti. (a) Lateral view of interior 
face of left valva; (b) Dorsal view of left valva. Top: Line drawing, prep. 1339, coll. 9 
July 1955. Bottom: Shade drawing, prep. 1341, coll. 13 July 1955. 


VOLUME 40, NUMBER 2 99 


Tegumen 














Fic. 2. Male Phoebis editha from Port-au-Prince, Haiti. (c) Lateral view of right side 
of genitalia (valvae, aedeagus, furca removed); (d) Lateral view of right side of furca; 
(e) Lateral view of right side of aedeagus; (f) Dorsolateral view of left side of distal end 
of aedeagus; (g) Dorsal view of uncus and tegumen. Top: Line drawing, prep. 1339. 
Bottom: Shade drawing, prep. 1341. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


100 














uorjoal 
Dy ipa DY} -o1d pajyutod yy 
‘qd ul se Buoy se yey pyypa *q ul -1pa ‘q Ul ueyy pez} py}1pa ‘qd ul ueyy pue peaimo A] 
uey} soul ATWYSI[G UeY} IoJoUTeIP Jesse] JO -O19TOs APY SI] S1OWY JaMoOIIeU yng ‘IasUOT -UdAZUN a8pa [eSIq saDUUAS ‘g 
anu 
aDUUaS *d Ul SB aDuuas ‘d ul ueuy -uas ‘d ul uey} pez aDuuas “d ul peAaimo 
BUC] SB 901M} JSOWY JaJBUIeIP 1938913 JO -O1ajos ATIAvY S1OJY UY} api ynq “Ie}10Y4S A[uaA9 aspa [eysiq  vyypad 
sayeue ovr[ided SI[eUISeAS Ue [Joe] Pu wIN319} YIg ue uIn319} 4g WINUSIS sotoeds 
JO aqoy [e1]UaA, uiIn319} y9g Aq pewio; suly Jor1eque sisAydody 
‘ODUUAS -d pue DY}Lpa ‘d jo eljeyues o[ewot u99M)}oq s90UsIO HIG G6 AITAV EL 
re ee SS 
DY} 
MOIA [eSIOP -1pa ‘qd Ul se apim UOISUd} Dyipa BA[BA JO UIS 
UI snoq[nq se d0IM], ‘yuIod -xa pojurod ‘qd ul se Zuo] DYpa “d -I8UI [SIP 
jou pus [e}sSIq ® 0} BUlIode} Ye] 7 yyw ‘xofdurop se jjey-2uO UI UPY} JOP uey} Iasu0'T apuuas ‘J 
apuuas ‘J ut 
MOIA se opin se J[ey-2uO aDu BABA jo uIs 
[es1op ul snoq ‘pus yeastp p9jutod aid -uas ‘qd UI SB aDuuas ‘d ul -Iew yeastp 
-[nq pus [eIsIq YM ‘eorpurfAD -UlIs ‘papunoy BUCO] SB 901M |, uey} IOMOLIVN uey} 1e,10Y§ py}1pa ‘d 
a a a ee Ee 
snouy) BARA JO ssad0id [es10g BA[eA Jo xody ssoooid Areyjndury snjnooes BATRA JO setoeds 


UIBIVUI [PSIOG 


a ee ee 
————————— 


‘apuuas ‘gq pue vYyIIpa ‘J jo eljeyues afew UseMjoq ssoUsIEHIqG ‘TT ATAVL 


VOLUME 40, NUMBER 2 101 


Ostium Bursae 


Antrum Tergum No.7 
Ductus Seminalis 
Appendix Tergum No.8 


REN SS 
Suprapapillary 


PrOGess 

Lamella Postvaginalis 
Tergum No.7 sss 

Tergum No.8 (abr Lamella Antevaginalis 

Apophysis Anterior @ |@2/)/ suprapapillary Process 


Papilla Analis 


Apophysis Anterior 


Ostium Bursae 
Sinus Vaginalis 


Antrum + 
Calyptra 


Flap 
Lamella Antevaginalis 





Fic. 3. Female Phoebis editha from Port-au-Prince, Haiti. (h) Dorsal view of geni- 
talia; (i) Lateral view of left side of genitalia (corpus bursae, ductus bursae, appendix 
bursae omitted). Top: Line drawing, prep. 1603, coll. 27 July 1955. Bottom: Shade draw- 
ing, prep. 1604, coll. 2 July 1954. 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


‘ | x 
* ae 


As SAS | 


LIOR 
Z AOE 
z hse <>. 








Fic. 4. Male Phoebis sennae from Haiti. (a) Lateral view of interior face of left 
valva; (b) Dorsal view of left valva. Top: Line drawing, prep. 1340, coll. Gros Morne, 2 
July 1954. Bottom: Shade drawing, prep. 1342, coll. Port-au-Prince, 13 July 1955. 


VOLUME 40, NUMBER 2 103 





1342 


Fic. 5. Male Phoebis sennae from Haiti. (c) Lateral view of right side of male 
genitalia (valvae, aedeagus, furca removed); (d) Lateral view of right side of furca; (e) 
Lateral view of right side of aedeagus; (f) Dorsolateral view of left side of distal end of 
aedeagus; (g) Dorsal view of uncus and tegumen. Top: Line drawing, prep. 1340. Bottom: 
Shade drawing, prep. 1342. 


104 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fic. 6. Female Phoebis sennae from Haiti. (h) Dorsal view of genitalia; (i) Lateral 
view of left side of genitalia (corpus bursae, ductus bursae, appendix bursae omitted). 
Top: Line drawing, prep. 1605, coll. Gros Morne, 4 July 1954. Bottom: Shade drawing, 
prep. 1606, coll. Port-au-Prince, 13 July 1955. 


VOLUME 40, NUMBER 2 105 


DESCRIPTION OF PHOEBIS EDITHA GENITALIA 


Male genitalia (Figs. 1 & 2). Valva flat; dorsal, proximal and distal margins concave; 
ventral margin convex; dorsal margin shorter than distal margin; distal margin about % 
length of ventral margin. Sacculus well defined, greatest width about ™% total length. 
Ampullary process cylindrical, sclerotized, gently curved toward proximal margin of 
valva, possessing a rounded apex, and covered with heavily sclerotized, stiff, cylindrical 
batons with bifurcate distal ends; overall length of ampullary process about % that of 
dorsal margin of valva. Apex of valva rounded; junction between dorsal and distal mar- 
gins of valva possessing a cylindrical, inward directed, heavily sclerotized dorsal process, 
about % length of dorsal margin of valva, with pointed distal end. 

Uncus fused with tegumen, without visible suture, possessing no definable proximal 
edge; distal part of uncus in lateral view tapering to a rounded point; uncus in dorsal 
view 1% times as wide as in lateral view, possessing blunt and imperceptibly bulbous 
apex. 

Tegumen forming shallow dome; peduncles poorly defined. Vinculum in lateral view 
about same length as combined length of tegumen and uncus. 

Saccus about % as long as combined length of tegumen and uncus, bent downward, 
with rounded distal end. 

Furca composed of two dorsal and two ventral processes, latter about % as wide and 
1% times as long as former. 

Aedeagus about 1% times as long as combined length of tegumen and uncus, bent 
upward along basal %, downward along distal %, possessing a single dorsal, flat spine 
near distal end, and a single ventrolateral flat spine basad of dorsal spine on left side, 
resulting in an asymmetrical arrangement; vesica with two oblong, sclerotized cornuti 
near distal end of aedeagus. 

Female genitalia (Fig. 3). Corpus bursae membranous, diaphanous, oblong, flask-shaped 
and expansible (thus of varying size); surface of membrane possessing numerous minute 
excrescences in the form of dots; a single oblong signum present near junction with 
ductus bursae; signum perpendicular to longitudinal axis of corpus bursae; dorsal and 
proximal part of signum possessing numerous spines. 

Appendix bursae likewise diaphanous and membranous, devoid of excrescences, spher- 
ical in shape; connected to corpus bursae by a diaphanous tube. 

Ductus bursae tubular, diaphanous; antrum sclerotized and about three times as long 
as ductus bursae; ductus seminalis arising dorsally from junction between ductus bursae 
and antrum. 

Lamella antevaginalis massive, sclerotized, shaped like a locomotive “cow catcher’; 
laterally fused with 8th tergum, forming with it a complete, uninterrupted ring with no 
visible suture; lamella postvaginalis with a movable protuberance, the calyptra, composed 
of lightly sclerotized and intricately folded membranes, blocking ostium bursae; ostium 
bursae laterally flanked by two free-standing membranous flaps. 

Apophyses anteriores of 8th tergum sclerotized; papillae anales bilobed; ventral lobe 
about half as wide as dorsal, but equal in length to it; dorsum of membranous area 
between 8th tergum and papillae anales with rounded, lightly sclerotized suprapapillary 
processes. 


DIFFERENCES BETWEEN GENITALIA OF 
PHOEBIS EDITHA AND PHOEBIS SENNAE 


Male genitalia (Figs. 1, 2, 4, 5). The differences are summarized in Table 1. 
Female genitalia (Figs. 3, 6). The differences are summarized in Table 2. 


The structural differences between P. editha and P. sennae, together 
with the fact that these butterflies are sympatric, synchronous, and 
have different larval foodplants, show that these taxa are specifically 


106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


distinct, and not, as some authors have suggested, forms of the same 
species. 


ACKNOWLEDGMENT 


I thank C. L. Remington for allowing me to borrow and dissect the genitalia described 
and figured here. 


LITERATURE CITED 


ABRERA D’, B. 1981. Butterflies of the neotropical region. Part 1. Papilionidae & Pier- 
idae. Lansdowne, East Melbourne. 172 pp. 

BUTLER, G. 1870. Descriptions of six new species of Callidryas. Trans. Entomol. Soc. 
Lond. 1870:9-12. 

Coutsis, J. G. 1983. Notes concerning certain West Indian butterflies. Entomol. Rec. 
95:113-114. 

Hiccins, L. G. 1975. The classification of European butterflies. Collins, London. 320 pp. 

RILEY, N. D. 1975. A field guide to the butterflies of the West Indies. Collins, London. 
224 pp. 

TUXEN, S. L. (Ed.) 1970. Taxonomist’s glossary of genitalia of insects. 2nd ed. Munks- 
gaard, Copenhagen. 359 pp. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 106 


GENERAL NOTE 


SMALL-NICOLAY COLLECTION TO SMITHSONIAN 


The National Museum of Natural History (Smithsonian Institution) is receiving the G. 
B. Small, Jr.—Col. S. S. Nicolay Collection of New World Butterflies. The scientific value 
of the collection is inestimable. It contains more than 3,800 species including about 450 
undescribed taxa. Its representation and identification of New World Hesperiidae, Lycae- 
nidae, and Riodinidae are now better than that in most museums. Coverage of Panama 
is approximately 98% complete, making its butterfly fauna better known than that of any 
other continental neotropical country. Geographically variable species, particularly from 
Panama, are represented by long series from many localities. Because so much neotropical 
forest has been destroyed, many of these specimens represent a unique record of the 
original fauna. Besides Panama, the collection is rich in material from Brazil, Costa Rica, 
Ecuador, Peru, and the United States. 

The Small—Nicolay Collection contains 98,500 specimens, of which more than 42,500 
are spread. There are 237 paratypes. 

Lepidopterists who expect to be in the Washington, D.C., area may visit the Smith- 
sonian and examine the collection by prearrangement. 


ROBERT K. ROBBINS AND J. F. GATES CLARKE, Department of Entomology, MRC 
NHB 127, National Museum of Natural History, Smithsonian Institution, Washington, 
D.C. 20560. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 107-123 


GENUS DIPTYCHOPHORA ZELLER AND A RELATED NEW 
GENUS STENEROMENE FROM THE NEOTROPICAL 
REGION (PYRALIDAE: CRAMBINAE) 


DAVID E. GASKIN 


Department of Zoology, University of Guelph, 
Guelph, Ontario, Canada NIG 2W1 


ABSTRACT. Diptychophora Zeller a Neotropical-southern Nearctic fringe genus of 
the subfamily Crambinae, is redefined. The type species D. kuhlweini Zeller, presently 
a junior synonym of D. azanalis (Walker), is resurrected and shown to be distinct. 
Diptychophora diasticta is described as new, D. subazanalis Bleszynski is elevated to 
specific rank, and Mysticomima desmoteria Meyrick is transferred to Diptychophora 
from its present position in Pyraustinae. Diptychophora azanalis is transferred, along 
with Pareromene nymphocharis (Meyrick), to the new genus Steneromene which is 
defined and distinguished from Diptychophora. 


The genus Diptychophora Zeller (Pyralidae: Crambinae: Diptycho- 
phorini) originally contained a single species from Brazil, D. kuhlweini 
Zeller, but subsequently became a repository for scores of small Cram- 
binae from all over the world except the northern Holarctic (Meyrick 
1931-88, Bleszynski & Collins 1962). Bleszynski (1965) determined 
that Pareromene Osthelder, erected for one species, P. rebeli, from 
Crete was the appropriate genus for all Old World “Diptychophora’’, 
and this name has since been used for these insects (Bleszynski 1966, 
1970, Gaskin 1971, 1974a, 1974b, 1975). However, there are problems 
relating to the continued use of this name (Gaskin 1985). Bleszynski 
(1967) formally redefined Diptychophora Zeller as an exclusively neo- 
tropical-southern Nearctic fringe genus with four species and one sub- 
species, but this is unsatisfactory for several reasons. His “‘subspecies’’ 
subazanalis deserves full specific rank within Diptychophora while 
azanalis requires a new genus to exclude it from Diptychophora. Ble- 
szynski also transferred the balance of named New World forms (then 
totalling nine species) to Pareromene. The status of these will be ex- 
amined elsewhere. 

During a long-term revision of the Diptychophorini of the world, 
all the above material was re-examined by the author and Michael 
Shaffer of the British Museum (Natural History), together with new 
finds from several collections not seen by Bleszynski. The purpose of 
the present paper is to redefine the genus Diptychophora, since the 
diagnosis given by Bleszynski (1967), essentially in three lines, is in- 
adequate; to resurrect the type species D. kuhlweini Zeller, since it is 
not synonymous with D. azanalis (Walker) as indicated by Bleszynski 
and Collins (1962) and Bleszynski (1967); to transfer one species de- 
scribed in Pyraustinae to Diptychophora; to describe a new species of 


108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the genus from Brazil; and to define a new genus Steneromene to 
contain D. azanalis (Walker) and Pareromene nymphocharis (Mey- 
rick). 

The following abbreviations for institutions are used in the text: 
BMNH (British Museum (Natural History)); CNIC (Canadian National 
Insect Collection, Ottawa); UMO (University Museum, Oxford, UK); 
USNM (United States National Museum of Natural History). In de- 
scriptions of male genitalia the LMB ratio refers to the length-median 
breadth ratio of the aedeagus. Decimals indicate the position of fea- 
tures, as a proportion of the total length of a structure or organ. In the 
forewing, measurements along the costa are taken from the base, those 
along the termen/margin from the apex, and those along fascia from 
the costa. In the male genitalia, measurements along the uncus, gna- 
thos, valva, valval costa and aedeagus are from the base of each. In 
the female genitalia, measurements along the ductus bursae are made 
from the ostium. 


Diptychophora Zeller 


Diptychophora Zeller (1866). Type species Diptychophora kuhlweini Zeller (1866) (by 
monotypy). 

Scissolia Barnes & McDunnough (1914). Type species Scissolia harlequinalis Barnes & 
McDunnough (1914) (by monotypy) [Syn. Bleszynski 1967]. 

Colimea Dyar (1925). Type species Colimea incisalis Dyar (1925) (by monotypy) [Syn. 
Bleszynski 1967]. 

Mysticomima Meyrick (1981). Type species Mysticomima desmoteria Meyrick (1931) 
(by monotypy) [New synonymy (from Pyraustinae)]. 


Revised description. In forewings, Sc and R, concurrent, R, vestigial or absent. In 
hindwings, M,, Cu, arising from common stalk, or individually from distal margin of 
hindwing cell. Male genitalia with characteristic “fish hook” gnathos, tubular, tapered 
to apex, hooking occurring at junction of basal elements. Saccus almost vestigial, juxta 
simple. Setulose valva highly modified, secondarily simplified, in most species broader 
than long, terminating distally in dorsal and ventral blunt lobes. Female genitalia simple: 
Antrum usually broad, membranous, ductus bursae also broad with or without subantral 
expansion. Corpus bursae bearing scobinate patch or distinct signum, usually crescentic. 


Key to Species of Diptychophora 


1 6: Sclerotized anellar structure present around aedeagus. [Fe- 
male net known) 03. sve OS diasticta 
- Selerotized anellar structure absent (eee 2 
2(1) 6: Ventral margin of valva slightly rounded, but with saccular 
region not developed into distinct lobe. ?: Ductus bursae 
with secondary swelling anterior to corpus bursae .... kuhlweini 
— 6: Ventral margin of valva terminating in a distinct project- 
ing, saccular lobe. ¢: Ductus bursae without secondary 
swelling anterior to corpus bursae 3 


VOLUME 40, NUMBER 2 109 


3(2) 6: Vinculum broad, half as wide as length of uncus; gnathos 
tapered to smoothly pointed apex. ?: Antrum a membra- 
nous funnel, wider than length of 8th abdominal tergum; 
corpus bursae with broad, diffuse scobinate patch, no dis- 
GINNe ESKO AUN relat aoe. lee AR Lasts Nivel asayig nes subazanalis 

— 6: Vinculum a narrow strip at base of valva: gnathos tapered 
abruptly, hooked dorsad. ¢: Antrum a membranous funnel 
if flared, considerably narrower than length of 8th tergum; 
corpus bursae with distinct sign oe 4 

4(3) 9: Ductus bursae lacking flared antrum, latter slightly 
“crimped”; 7th sternum with tapered and rounded poste- 
nommarczin. (Male not known] 225080 te incisalis 

— 6: as described in 3-. 2: Antrum of ductus bursae flared at 
entrance; 7th sternum not tapered and barely rounded pos- 
(ESTEAOPEN a gat = ETT dae oe en ee ee a 5 

5(4) 6: Costal lobe of valva relatively broad, length to width ratio 
about 2.2:1. 2: Antrum of ductus bursae membranous, with 
smooth margin; corpus bursae with small, crescentic nar- 
Low eanonizontalisionum: Be desmoteria 

— 6: Costal lobe of valva quite elongate, length to width ratio 
about 3.5:1. 2: Antrum of ductus bursae with slightly cren- 
ulate margin, somewhat sclerotized; corpus bursae with 
large, tapered, horizontal signum with infolded lower (an- 
CSPI MADAIRSTNAS U7 a ennai oo oc ee aa harlequinalis 


Diptychophora diasticta, new species 


Exterior description (Fig. 5). Alar expanse 12 mm (N = 1). Labial palpi, head, an- 
tennae brown, head with postcephalic, buff scale tuft. Thorax and abdomen brown with 
scattered darker scaling. Ground color of forewings creamy white, heavily irrorated with 
brown clouding. Basal fascia obsolete, position marked with scattered brown scales. An- 
temedial fascia cream, zigzagged, broad, thickly bordered with dark brown. Discal region 
clouded with brown, cream zone just distal of median transverse line. Costa with orange 
area at about 0.7 and small oblique central white bar. Postmedial fascia wider toward 
dorsum, also becoming less distinct, zigzagged from 0.5, with thick dark brown margins. 
Terminal zone cream, heavily clouded with brown. Margin with stripe of bright orange 
from 0.4—0.8, and row of 4 black spots. Apical zone clearly delineated by narrow white 
line extending from 0.8 of costa to subapical identation on termen, orange brown proxi- 
mally, white distally, wedge of yellow scales on costal extremity. Cilia brown with darker 
brown apices. Hindwings dark brown. Ventral surfaces dark brown, apical and terminal 
markings repeated from dorsal surface. 

Male genitalia (Fig. 9) (N = 1). Uncus simple, broad, spatulate, apically rounded, slight 
subapical constriction. Gnathos triangular in cross section, slightly tapered to apex, angled 
sharply dorsad at about 0.4. Tegumen simple, vinculum a strip at base of valva, saccus 
small, pyramidal. Juxta an elongate, apically tapered plate; hooked, double transtillalike 
structure present above juxta. Valva about 2.2x length of uncus, half as broad as long, 
broad posteroventral lobe. Sacculus, costal region of valva undeveloped. Aedeagus tu- 
bular, truncate, curved slightly dorsad, huge relative to rest of genitalia, about 1.8x 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in 


ee es 
() 


xe 


eee 
P=} 
= 
aS 
a 
a 





Fics. 1-8. Forewings of Steneromene and Diptychophora species. 1, Steneromene 
azanalis; 2, S. nymphocharis; 3, Diptychophora kuhlweini; 4, D. subazanalis; 5, D. 
diasticta; 6, D. desmoteria; 7, D. harlequinalis; 8, D. incisalis. 


length of valva. LMB ratio about 10:1, cornuti absent, scobinate patch of vesica in 


subapical region. 
Type. Holotype 6. Prov. de Bahia, Sao Antonia de Barra, Brazil, II. XII 1888, Gounelle, 


BMNH, genit. prep. Pyralidae 15351. 


Discussion. Known only from the holotype, which is from eastern 
Brazil. Except for the development of a sclerotized, rather than mem- 
branous transtilla, the male genitalia are typical of the genus. 


Diptychophora kuhlweini Zeller 


Diptychophora kuhlweini Zeller (1866). [Holotype not located despite search of collec- 
tions in Britain and Europe by author, M. Shaffer, and other curators. ] 


VOLUME 40, NUMBER 2 ABS 


Diptychophora kuhlweini Zeller; Bleszynski & Collins (1962); Bleszynski (1967); erro- 
neously synonymized with D. azanalis (Walker). 


Exterior description (Fig. 3). Alar expanse 12 mm (N = 4). Labial palpi whitish, some 
dark scaling; head, thorax, largely dull white. Legs, straw; abdomen white, scattered dark 
scaling. Ground color of forewings shining white. Basal and antemedial fasciae nearly 
straight, proximally bright yellow ochre, distally blackish brown. Postmedial fascia proxi- 
mally broad and blackish brown, distally broad and ochreous yellow. Subterminal line 
turning at right angles toward termen at about 0.2, touching margin at primary inden- 
tation of termen. Terminal zone white, a few yellowish scales near margin, two black 
spots at 0.6 & 0.7. Apical zone delimited by marginal indentation (a secondary inden- 
tation occurs at nearly 0.5) and by thin silvery white line running from costa at about 
0.8 to angle of subterminal line, also joining with a weaker thin line angling proximally 
from the costa. Apical zone bright ochreous yellow, white wedge of scales at extremity, 
a horizontal, white, pendant-shaped mark above indentation. Hindwings shining white, 
few dark scales near apex. 

Male genitalia (Fig. 10) (N = 3). Uncus broad, tapered, spatulate, apically rounded. 
Gnathos triangular in cross section, flat surface dorsad, sharply pointed, curved or angled 
very acutely dorsad at about 0.4. Tegumen simple, sclerotized bandlike margins. Vin- 
culum a narrow strip at base of each valva. Saccus very small, juxta a weak, oval plate, 
almost membranous. Valva trapezoid in shape, basal length barely greater than dorso- 
ventral width, dorsal margin about 1.8 width; dorsal margin sclerotized, apex tapering, 
rounded. Aedeagus small, barely 0.5 valva, tubular, apically truncate, LMB ratio about 
5.0:1. Cornuti absent. 

Female genitalia (Fig. 14) (N = 1). Anal papillae triangular, moderately sclerotized, 
weakly fused; 8th abdominal tergum narrow; anterior apophyses short, nearly as long as 
posterior apophyses; 7th abdominal sternum unmodified, slightly rounded posteriorly. 
Antrum weak, membranous, masked by margin of 7th sternum. Ductus bursae 6-7 x 
length of posterior apophyses, membranous, flattened to about 0.5, where it is swollen 
and lightly reticulate. Corpus bursae with two crescentic signa. 

Material examined. BRAZIL: 1 6, Castro, Parana, 950 m, Jones, no date BMNH, genit. 
prep. Pyral. 15358; 1 6, Petropolis, Prov. Rio de Janeiro, no date, BMNH, GS-5083-SB; 
1 6, Nova Teutonia, —.X.1948, F. Plaumann, BMNH, Pyral. 16825; 1 9, Guaraquefaba, 
P.R., 7.XII.1970, V. O. Becker Collection, Brasilia, spec. no. 11,428. 


Discussion. This species is probably widely distributed across the 
southern part of the Brazilian shield although its specific habitat is 
unknown. The flight period includes at least October and December. 

Bleszynski (1967) noted that “too little of the typical azanalis ma- 
terial is available . . . to decide whether suwbazanalis is a distinct species 
or only a subspecies’. It would appear that unknowingly, Bleszynski 
never examined any true azanalis material at all, since M. Shaffer and 
I were only able to locate the type and one other specimen, both in 
UMO. Had Bleszynski seen the genitalia of the type slide he would not 
have made the remark quoted above because congenerity is out of the 
question. M. Shaffer first drew my attention to inconsistencies in the 
forewing characters within the “azanalis’” and “subazanalis’ series 
accumulated by Bleszynski and stored at BMNH. When the specimens 
with the thicker, solid postmedial fascia angled beneath the apex were 
compared to the illustration of kuhlweini given by Zeller (1866), they 
matched exactly. We then compared the genitalia between specimens 
segregated on the basis of forewing pattern and again found consistent 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


differences, much more marked in the female than in the male (Figs. 
9, 11, 14-16). We concluded that kuhlweini was a valid species of 
Diptychophora, and that synonymization with azanalis was invalid, 
and, furthermore, that swbazanalis should be elevated to full specific 
rank within the genus. Both known specimens of true azanalis (Fig. 
1) were also noted to differ consistently in forewing characters from 
subazanalis (Fig. 4), the postmedial fascia of the former being thin, 
single and curved towards the costal margin. The genitalia of both 
sexes of azanalis (Figs. 20, 22) are unlike those of any Diptychophora. 


Diptychophora subazanalis Bleszynski, new status 


Diptychophora azanalis subazanalis Bleszynski (1967). [¢ genitalia, mislabelled D. ex- 
aminalis subexaminalis. | 


Exterior description (Fig. 4). Alar expanse 8-12 mm (N = 23). Labial palpi, head, 
white with yellow, brown; thorax shining white, yellow lappets, chocolate brown shoul- 
ders. Ground color of forewings white. Basal and antemedial fascia broad, nearly straight, 
each proximally bright yellow, distally chocolate brown. Postmedial fascia narrow, com- 
posed of pair of slender dark brown parallel lines, angled sharply toward termen at about 
0.2, angled again at about 0.6. In Peruvian specimens, second angle of inner transverse 
line sometimes detached as a dark brown blotch. A broad strip of bright yellow extends 
along proximal side of inner line. Apical zone bright ochreous yellow; a white streak 
curves through it from costa to termen. Terminal region filled anteriorly with yellow, 
posteriorly with white; margin bears two black spots at 0.5 and 0.7. Cilia brown with 
yellow clouding. Hindwings and their cilia shining white. Ventral surfaces brown on 
forewings, white on hindwings. 

Male genitalia (Fig. 11) (N =3). Uncus simple, broad, apically rounded, spatulate. 
Gnathos tubular, tapering sharply to point, curved acutely dorsad at about 0.4-0.5. Tegu- 
men broad, divided into wide dorsal and ventral bands, juxta a simple oval plate, vin- 
culum broad, half as wide in profile as length of uncus, saccus almost negligible. Valva 
characteristically wider than long, drawn into blunt, double apex dorsally and ventrally; 
costa of valva represented by thin sclerotized margin, sacculus undeveloped. Aedeagus 
short, tubular, apically truncate, LMB ratio about 6:1, cornuti absent. 

Female genitalia (Figs. 15, 16) (N = 4). Anal papillae broad, slightly sclerotized mar- 
ginally, relatively short posterior apophyses about 1.5 length of papillae; 8th abdominal 
tergum and anterior apophyses each about half length of posterior apophyses, 7th ab- 
dominal sternum unmodified. Antrum a very broad, shallow, basally constricted, weak 
funnel, ductus bursae otherwise broad, about 4.5 length of posterior apophyses. Ductus 
seminalis joining at about 0.4-0.5. Corpus bursae with huge, crescentic scobinate area or 
diffuse signum. 

Types. Holotype 4, SURINAM. Zanderij, Boven, Para district, 25.IV.1927, Cornell 
University collection, genit. prep. GS-5140-SB. 

Paratypes, SURINAM: 2 4, 2 2, as above but 19, 20.I1V.1927 in case of 2, Cornell 1 4, 
CNIC, Ottawa, 1 ?; BMNH, 1 4, 1 2; genit. preps. GS-5141-SB 2, GS-4383-SB °. GUYANA: 
2 6, Tumatumari, Potaro River, 27-29.VI.1927, Cornell, genit. prep. Cornell slide no. 2 
(M. Shaffer prep.). 

Other material examined. GUYANA: 7 6, Atkinson airfield, nr. Georgetown, 1955, 
Lyall, BMNH (all lack abdomens); 1 6, Bartica, -.1.13, BMNH (no abdomen); 1 6, same 
locality, 6.V.1901, BMNH; 1 2, same locality BMNH, Pyral. 15355 ¢. PERU: 2 4, 1 9, 
Iquitos, -.V.1920, Parish, BMNH, Pyral. 15354 (1 6 without abdomen). BRAZIL: 1 9, 
Para (=Recife), Prov. Pernambuco, Serra de Communaty, 1.]I.1893, Gounelle, BMNH; 
1 6, 1 8, Amazon Reserva Ducke, km, 26, Manaus-Itacoatiara Highway, 21.IV.1971, E. 
G. Munro, CNIC, Ottawa. 


VOLUME 40, NUMBER 2 118 










W 
I) 


) SS ep 
we es Teh 


ep tO MO nq Ses ~- 
Start ene, ye rae 


y 


Ie 


Fics. 9, 10. Male genitalia of Diptychophora. 9, D. diasticta holotype, aedeagus 
(left), transtilla in lateral aspect (center), posterior aspect of genitalia with left valva 
(right); 10, D. kuhlweini, aedeagus (left), genitalia with left valva (right). Scale = 1 mm. 


Discussion. The male and female genitalia are redescribed here. In 
the case of the male, Bleszynski’s brief comments are ambiguous; he 
overlooked some structures in his drawing, and the proportions are 
misleading. His drawing of the female was made from a poor prepa- 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 11, 12. Male genitalia of Diptychophora. 11, D. subazanalis, aedeagus (left), 
posterior aspect of genitalia with left valva (right); 12, D. desmoteria lectotype, aedeagus 
(left), genitalia with left valva (right). Scale = 1 mm. 


ration. The amount of material collected for this species is so great 
compared to some others that it must probably be either one of the 
commonest Diptychophora species, or perhaps particularly attracted 
to light. It has been taken in the Amazon Basin from Iquitos to Para, 


VOLUME 40, NUMBER 2 DS 





Fic. 18. Male genitalia of Diptychophora harlequinalis, aedeagus (left), posterior 
aspect of genitalia with left valva (right). Scale = 1 mm. 


N to the Caribbean coast and S to the edge of the Brazilian Shield; the 
flight period is known to include January—June. Explanation of eleva- 
tion of subazanalis to full specific status appears in the discussion of 
D. kuhlweini. 


Diptychophora incisalis (Dyar) 


Colimea incisalis Dyar (1925). 
Scissolia incisalis (Dyar); Bkeszynski (1966). 
Diptychophora incisalis (Dyar); Bleszynski (1967). 


Exterior description (Fig. 8). Alar expanse (?) 14-15 mm (N = 2). Details of external 
characteristics were adequately provided by Dyar (1925). The male is unknown. 

Female genitalia (Fig. 17) (N=1). Anal papillae weakly fused, small, about 0.5 x 
length of posterior apophyses, 8th tergum and anterior apophyses shattered in Bleszyn- 
ski's paralectotype preparation, 7th sternum tapered, rounded posteriorly. Antrum a 
weak funnel, ductus bursae about 6x length of posterior apophyses, ductus seminalis 
joining at about 0.3. Corpus bursae with single, huge strong crescentic signum, basal 
margin sharply turned introrse into bursae. 

Types. Lectotype and paralectotype 2, MEXICO, Colima, -.VIII.1923, Muller, type 
27503, USNM, genit. prep. GS-6117-SB. 


Discussion. The female genitalia are redescribed because Bleszyn- 


116 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fics. 14-16. Female genitalia of Diptychophora, ventral aspect. 14, D. kuhlweini; 
15, D. subazanalis paratype; 16, D. subazanalis variation in form of signum. Scale = 1 
mm. 


ski’s diagnosis and illustration are ambiguous. This highly patterned 
insect has so far been reported only from the type locality in western 
Mexico during August. The lectotypic series was selected by S. Ble- 
szynski (1966). 


Diptychophora desmoteria (Meyrick), new combination 


Mysticomima desmoteria Meyrick (1931) (from Pyraustinae). 


Exterior description (Fig. 6). Alar expanse 14-15 mm (N = 6). External characteristics 
were described in detail by Meyrick (1931). 

Male genitalia (Fig. 12) (N = 2). Uncus simple, broad, apically rounded, spatulate, 
slightly constricted subapically. Gnathos triangular in cross section, curved sharply dorsad 
at 0.2, only slightly tapered until near the blunt apex, which is angled dorsad. Tegumen 
simple; juxta a suboval, relatively weak plate; saccus simple, small, rounded apically; 
vinculum a narrow band at valval base. Valva about 3x length of uncus, 0.8 as broad 


VOLUME 40, NUMBER 2 BET 





Fics. 17-19. Female genitalia of Diptychophora, ventral aspect. 17, D. incisalis 
paralectotype, 17a, signum in lateral aspect; 18, D. desmoteria; 19, D. harlequinalis, 
19a, signum in lateral aspect. 


as long, bifurcating into dorsal and ventral lobes, latter moderately pointed. Aedeagus 
about 1.2x length of simple, tubular, apically expanded valva, LMB ratio about 7.5- 
8: 1. No cornuti. 

Female genitalia (Fig. 18) (N=1). Anal papillae small, weak; posterior apophyses 
about 3x length of papillae; 8th tergum narrow, about 0.3x length of posterior apo- 
physes, anterior apophyses about 0.5 posteriors; 7th sternum unmodified. Antrum a 
weak, simple funnel. Ductus bursae about 3.5 length of posterior apophyses, ductus 
seminalis joining at about 0.3. Corpus bursae with single, narrow, crescentic signum. 

Types. Lectotype 6, COSTA RICA: San Jose, 1922, BMNH, genit. prep. Pyral. 15096. 
Selected by author and M. Shaffer (BMNH) and designated here. Paralectotypes, COSTA 
RICA: 3 4, 1 9, data as above, BMNH, genit. preps. Pyral. 2475 4, 15096 2. 

Other material examined. 1 6, COSTA RICA: 3.11.24, Schmidt, BMNH. 


Discussion. While curating BMNH Pyralidae, M. Shaffer noticed 
that desmoteria possessed not only a strong superficial resemblance in 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


both pattern and color to Diptychophora harlequinalis, but also the 
cleft forewing apex typical of most Diptychophorini. Dissection of the 
genitalia confirmed immediately that the species belonged in this ge- 
nus, not in the Pyraustinae. 


Diptychophora harlequinalis Barnes & McDunnough 


Scissolia harlequinalis Barnes & McDunnough (1914). 
Scissolia harlequinalis Barnes & McDunnough; Bleszynski (1966). 
Diptychophora harlequinalis (Barnes & McDunnough); Bleszynski (1967). 


Exterior description (Fig. 7). Alar expanse 15 mm (N = 6). External features described 
by Barnes and McDunnough (1914). 

Male genitalia (Fig. 13) (N = 2). Uncus simple, broad, spatulate, apically rounded; 
gnathos triangular in cross section, tapered, curved sharply dorsad at about 0.4, apically 
pointed, point slightly turned dorsad. Tegumen simple, strong ventral margins; juxta a 
squat, semicircular, moderately sclerotized simple plate; vinculum a narrow strip at base 
of valva; saccus simple, small, rounded anteriorly. Valva about 3.5x length of uncus, 
broadly bifurcate distad, lacking costal or saccular development. Aedeagus tubular, api- 
cally truncate, slightly curved dorsad, equal to valva in length, LMB ratio 89:1, cornuti 
absent. 

Female genitalia (Fig. 19) (N = 3): Anal papillae weakly fused, marginally sclerotized, 
posterior apophyses about 1.25 length of papillae; 8th tergum about 0.9x length of 
posterior apophyses; anterior apophyses present only as short, broad, pointed prongs less 
than length of tergum; 7th abdominal sternum unmodified. Antrum weak, flared funnel 
with slightly crenulate margin; ductus bursae about 6x length of posterior apophyses; 
ductus seminalis joining at about 0.3. Corpus bursae with single crescentic signum. 

Types. Holotype 6, U.S.A., Arizona, no date, Cornell collection. 

Other material examined: USA: 1 6, Arizona, Oslar, Huachuca Mts., 28. VIII.1903, 
BMNH, genit. prep. Pyral. 15095; 1 9, Arizona, Madera Canyon, Santa Rita Mts., 
25. VIII.1946, Comstock & Martin, Cornell, genit. prep. Cornell #2 (M. Shaffer prep.); 1 
6, same locality, 25.VIII.i946, CNIC, Ottawa; 2 9, same locality, 27.VII.1947 and 
3. VIII.1959, R. W. Hodges, genit. prep. EGM 1744 CNIC, Ottawa, genit. prep. EGM 
1745 and 4912-SB. 


Discussion. This bright yellow and black little moth has yet to be 
recorded outside Arizona, where its habitat is montane forest. 


Steneromene, new genus 


Type species ?Zebronia azanalis Walker (1859). 


Description. Hindwing venation characterized by M, and Cu, free as in most Old 
World Diptychophorini, not stalked as in Diptychophora. Male genitalia characterized 
by elongate valva, definite apical lobate expansion (but not as extreme as in most species 
of Diptychophora), no development of costal region. Juxta bearing two pairs of apical 
horns. Saccus, vinculum exceedingly narrow. Female genitalia with eighth tergum re- 
duced and narrow, anal papillae fused dorsally, ductus bursae bearing spinose, sclerotized 
subantral globate sac, ductus seminalis arising from its dorsoposterior surface, or a short 
distance anterior to it. 


Steneromene azanalis (Walker), new combination 


?Zebronia azanalis Walker (1859). 
Usopteryx parvalis Walker (1865) [Erroneous synonymy by Bleszynski & Collins (1962).] 


VOLUME 40, NUMBER 2 119 





Fics. 20, 21. Male genitalia of Steneromene. 20, S. azanalis, aedeagus (left), pos- 
terior aspect of genitalia with left valva (right); 21, S. nymphocharis, paralectotype, 
aedeagus (left), posterior aspect with left valva (right). 


[Diptychophora kuhlweini Zeller (1866). [Erroneous synonymy by Bleszynski & Collins 
(1962). ] 

Diptychophora azanalis (Walker); Bleszynski & Collins (1962); Bleszynski (1967). 

[Diptychophora kuhlweini Zeller; Bleszynski (1967). [Erroneous synonymy. ] 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 22-24. Female genitalia of Steneromene. 22, S. azanalis holotype, ventral 
aspect; 23, S. azanalis, diagram of antrum in lateral aspect, a, ductus bursae; b, lamella 
postvaginalis; c, lamella antevaginalis; d, antral “pocket”; e, 7th sternum; f, antrum; g, 
subantral expansion; 24, S. nymphocharis lectotype, ventral aspect, a, diagram of an- 
trum, ventral aspect. 


Exterior description. Alar expanse 15 mm (N = 2) (Fig. 1). Labial palpi, head, thorax, 
silvery white, some brown scaling. Ground color of forewings shining white, some brown 
scaling along costa. Basal fascia reduced to short black bar on costa near wing base. 
Antemedial fascia mid-brown, lateral orange shading. Postmedial fascia mid-brown, very 
narrow, single, virtually straight from tornus to 0.2 from costa, where it curves smoothly 
inwards to meet costa. Terminal zone white, except for marginal brown line with some 
ochre shading from 0.4—0.6, within which are two black dots. Cilia brown. Apical zone 


VOLUME 40, NUMBER 2 12] 


orange ochre, with small white wedge of scales at costal extremity, and a large, irregularly 
shaped shining white central area, touching costa at one point, and caudally edged with 
black. Hindwings white, with pale brown cilia. Ventral surface pale brown, apical mark- 
ings of forewing repeated from dorsal surface. 

Male genitalia (Fig. 20) (N =1). Uncus, gnathos simple, tapered, bluntly pointed. 
Tegumen weak except for strong posterior margins. Vinculum narrow, about 0.3 as wide 
as uncus is long. Saccus broad, straplike, hardly developed. Juxta strong, massive, with 
two pairs of apical horns, outer pair serrate on their inner margins. Valva about 2.8 x 
length of uncus, striking concavity on inner surface from 0.2 to 0.4, costa strong but not 
developed into protrusions. Sacculus developed, apical prong at about 0.7 from base of 
valva. Aedeagus tubular, about 0.8 length of valva, cornuti absent, LMB ratio about 
8:1: 

Female genitalia (Figs. 22, 23a—g) (N = 1). Anal papillae broad, marginally sclerotized, 
about half as long as posterior apophyses. Anterior apophyses absent, 8th tergum about 
0.4x length of posterior apophyses, 7th sternum not tapered posteriorly, but posterior 
margin in-turned to form complex lodicular structure with lamella antevaginalis, struc- 
turally a broad, heavily sclerotized ventral collar to the antrum. Lamella postvaginalis 
strong, suboval, fused laterally with margins of antevaginalis, forming plate dorsal to 
small and narrow ostium. Both lamellae spinose and scobinate. Ductus bursae with scler- 
otized swelling immediately below antrum proper, bulging dorsad, containing a half 
reverse loop of ductus bursae, the latter about 5.5-6x length of posterior apophyses, 
broad, weak below subantral region, ductus seminalis joining at about 0.3. Corpus bursae 
with single small, circular signum. 

Type. Holotype 2, BRAZIL: “Rio”, “87”, UMO, type slide 1193. 

Other specimen examined. BRAZIL: “Rio”, 1 6 no other data, M. Shaffer genit. prep. 
1979/7 (UMO). 


Discussion. I have no information on the distribution of this species; 
“Rio” presumably refers to Rio de Janeiro. The juxta is proportionately 
large and sclerotized, and there is some sclerotization of the saccular 
region of the male valva. 


Steneromene nymphocharis (Meyrick), new combination 


Diptychophora nymphocharis Meyrick (1932). 
Diptychophora nymphocharis Meyrick; Bleszynski & Collins (1962). 
Pareromene nymphocharis (Meyrick) Bleszynski (1967). 


Exterior description (Fig. 2). External characters were described by Meyrick (1932). 

Male genitalia (Fig. 21) (N = 1). Uncus simple, tapered, curved slightly ventrad, apex 
blunt and slightly expanded. Gnathos broad, spatulate, apically rounded with scobinate 
margin, membranous except for margins. Tegumen reduced, weak, prominent dorsal 
and ventral straplike margins. Vinculum a narrow straplike structure at base of valva. 
Saccus short, rounded with ventromedial strengthening. Juxta an oval plate bearing two 
pairs of apical horns, probably anellar in origin but firmly fused with rest of plate. Inner 
pair twice length of outer pair. Valva long, broad, massive in comparison to rest of 
genitalia, about 4.5x length of uncus; apically expanded, with beginnings of saccular 
fold. Costa distinct to about 0.8 on dorsal margin, without protuberances. Lateral basal 
coremata present. Aedeagus long, 1.1 x length of valva, simple, tubular, LMB ratio about 
12:1, cornuti absent. 

Female genitalia (Fig. 24) (N =1). Anal papillae strongly fused in dorsal midline; 
anterior apophyses about 0.7 x length of posterior apophyses, 8th tergum less than half 
as long as anterior apophyses, 7th sternum rounded posteriorly, otherwise unmodified. 
Antrum a strong, scobinate, flared funnel. Lamella postvaginalis forming roof of antrum, 
but medially cleft. Ductus bursae about 3.5 x length of posterior apophyses, ductus sem- 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


inalis joining at 0.3, and arising from swollen, scobinate, subantral sac. Corpus bursae 
bearing single, small signum. 

Types. Lectotype 2, designated here, ARGENTINA: Alta Gracia, .32, “C.B.”, BMNH, 
genit. prep. Pyral. 15099. Paralectotypes, 1 6, 1 2, same locality but dated “2.34”, BMNH, 
Pyral. 15100 6. 


Discussion. Presently known only from one locality in Argentina 
(Sierra de Cordoba). As in azanalis, the male valva is not as strikingly 
modified from the elongate quadrate form typical of most Diptycho- 
phorini as in Diptychophora itself. Sclerotization of the antrum of the 
female ductus bursae is not as marked as in azanalis. 


ACKNOWLEDGMENTS 


The author thanks the Department of Entomology, British Museum (Natural History) 
for providing space, facilities and assistance during this study, and the Biosystematics 
Research Institute, Ottawa, the University Museum of Oxford, the Cornell University 
Museum, and the United States National Museum for loans of specimens including types. 
I especially thank Michael Shaffer at the British Museum for obtaining loans, dissecting 
type material, and offering his experience and advice at all stages of the work. 


LITERATURE CITED 


BARNES, W. & J. H. MCDUNNOUGH. 1914. Some new North American Pyraustinae. 
Cont. Nat. Hist. Lepid. N. Am. 2:223-246. 

BLESZYNSKI, S. 1965. Crambinae: 2 vols. text and plates. Vol. 1 of “Microlepidoptera 
Palaearctica”, Amsel, H. G., F. Gregor & H. Reisser (eds.), i-xlvii; 1-552; Pl. 1-183 
(1-31 col.). 

1966. Studies on the Crambinae (Lepidoptera). Part 43. Further taxonomic 

notes on some tropical species. Acta Zool. Cracov. 11:451-494. 

1967. Studies on the Crambinae (Lepidoptera). Part 44. New Neotropical gen- 

era and species. Preliminary checklist of Neotropical Crambinae. Acta Zool. Cracov. 

12:39-110. 

1970. New genera and species of tropical Crambinae (Studies on the Crambi- 
nae, Lepidoptera, Pyralidae, Part 48). Tijdschr. Ent. 113:1-26. 

BYESZYNSKI, S. & R. J. COLLINS. 1962. A short catalogue of the world species of the 
Family Crambidae (Lepidoptera). Acta Zool. Cracov. 7:197-389. 

Dyar, H. G. 1925. Some new American moths (Lepidoptera). Insec. Inscit. Menstr. 13: 
1-19. 

GASKIN, D. E. 1971. A revision of New Zealand Diptychophorini (Lepidoptera: Pyrali- 
dae: Crambinae). New Zeal. J. Sci. 14:759-809. 

1974a. The species of Pareromene Osthelder (Pyralidae: Crambinae: Diptycho- 

phorini) from the western South Pacific, with further notes on the New Zealand 

species. J. Entomol. (B) 43:159-184. 

1974b. The species of Pareromene Osthelder (Pyralidae: Crambini: Diptycho- 

phorini) from Malaysia, Indonesia and New Guinea. J. Entomol. (B) 43:185—208. 

1975. A revision of the Australian species of Pareromene (Lepidoptera: Pyrali- 

dae: Crambinae: Diptychophorini). Aust. J. Zool. 23:123-147. 

1985. Morphology and reclassification of the Australasian, Melanesian and Poly- 
nesian Glaucocharis (Lepidoptera: Crambinae: Diptychophorini). Aust. J. Zool. Suppl. 
ser. 115:1-75. 

MEyRICK, E. 1931-33. Exotic Microlepidoptera 4:33-192; 193-252; 253-448. 

OSTHELDER, L. 1941. Beitrag zur Kleinschmetterlinges fauna Kretas. Mitt. Miinch. 
Entomol. Ges. 31:365-70. 























VOLUME 40, NUMBER 2 123 


WALKER, F. 1859. List of the specimens of lepidopterous insects in the collection of 
the British Museum, London, 19:967. 

ZELLER, P. C. 1866. Beschreibung einiger amerikanischen Wickler und Crambiden. 
Entomol. Ztg., Stettin 27:137-157. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 123 


BOOK REVIEW 


BUTTERFLIES OF EUROPE, vol. 1. CONCISE BIBLIOGRAPHY OF EUROPEAN BUTTERFLIES, 
by Otakar Kudrna. 1985. AULA-Verlag GmbH, Postfach 1366: D-6200 Wiesbaden, West 
Germany. 447 pp. Octavo, hard bound. $67.00 (including shipping). 


This is the first in a series of planned volumes reappraising knowledge of European 
butterflies. Future volumes will cover all butterfly families except skippers; also, intro- 
duction to lepidopterology, ecology, and conservation. When completed, the series prom- 
ises to be the most in-depth study of European butterflies, and will include taxonomy, 
life histories, biogeography, ecology, behavior, etc. There is a minimum of typographical 
errors in this first volume. 

The bibliography lists about 6,000 references, primarily those between 1901 and 1983, 
the period before 1900 already having been covered by Horn and Schenkling, Derksen 
and Scheidung, Bang-Haas, Bretherton, Junk, and Kusnezov. The succinct Preface and 
Introduction discuss methods, purposes, history, 19th century works, and acknowledge- 
ments. Information content on ecology, biogeography, conservation, and taxonomic re- 
vision proves sparse for Europe. “The present bibliography is ... aimed to serve the 
needs of all students of the butterflies of Europe, whether they take primary or secondary 
interest in lepidopterology, whether they are professionals or amateurs, whether they are 
beginning students, or experienced scientists and/or university lecturers.” Kudrna esti- 
mates that over 50,000 titles bear directly or indirectly on the butterflies of Europe, 
with the included 6,000 being selected on merit and usefulness. He plans to update the 
bibliography with supplements, and eventually produce a larger, more comprehensive 
work of 10-15,000 references. Especially relevant to North American workers are the 
many general studies cited; part of our nearctic butterfly fauna was derived from the 
Palaearctic. 

This bibliography is a useful reference source that must have been a Herculean effort 
to prepare. Over 80% of the references were checked against the original works for 
citation accuracy. Europe presents special problems because of its multitude of languages 
and the scattered references. The completion of the eight volumes of the Butterflies of 
Europe toward the end of this century, produced by a team of modern specialists and 
edited by Kudrna, promises to be a major advance in the scientific study of butterflies. 


OAKLEY SHIELDS, 4890 Old Highway, Mariposa, California 95338. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 124-126 


GENERAL NOTES 


OBSERVATIONS ON THE DIURNAL GREGARIOUS ROOSTING OF 
OCALARIA SP. (NOCTUIDAE) IN COSTA RICA 


Gregarious roosting behavior (quiescent aggregations for sleeping or passing unfavor- 
able periods) has been documented for various diurnally active Lepidoptera (DeVries et 
al., Zool. J. Linn. Soc., in press). Among nocturnally active Lepidoptera, however, little 
evidence of a consistent roosting habit has been found. An exception is the well-docu- 
mented aestivation behavior of the Australian bogong moth, Agrotis infusa (Boisduval) 
(Noctuidae). During the summer when its larval host plants are not available, adults mi- 
grate to granite outcrops and there form dense aggregations (Common 1954, Aust. J. 
Zool. 2:223-263) reminiscent of those formed by the North American monarch butterfly, 
Danaus plexippus (Linnaeus) (Nymphalidae). Another type of diurnal roosting in a 
nocturnally active species is that of the skipper butterfly, Celaenorrhinus fritzgaertneri 
(Baily) (Hesperiidae). It forms roosts in caves and hollows during the dry season in Costa 
Rica, apparently as a mechanism for passing a time of year when conditions for larval 
development are unfavorable (DeVries et al., cited above). In contrast to the bogong 
moth, however, C. fritzgaertneri adults leave and return to the roost following a syn- 
chronous circadian rhythm. 

The genus Ocalaria (Noctuidae) was established by Schaus (1906, Proc. U.S. Nat. Mus. 
30:132) for a group of moths whose phylogenetic affinities are uncertain, but which is at 
present tentatively placed at the end of the subfamily Ophiderinae (Erebinae of authors) 
in thé U.S. National Museum of Natural History (NMNH) (R. W. Poole, pers. comm. ). 
The Ocalaria species treated here is a small, mottled gray-brown moth with an eyespot 
on both upper and lower surfaces of the forewing and a wingspan of about 1.5 cm (Fig. 
1). Based on NMNH collections, this undescribed species has been collected most often 
in tree buttresses in lowland areas of Panama (R. W. Poole, pers. comm.). We describe 
here a roost of the species from Costa Rica, and provide notes on its roosting behavior. 

In July and August 1983, a diurnal roost of Ocalaria sp. was observed on six separate 
occasions near the Sirena Station of Parque Nacional Corcovado in southwestern Costa 
Rica. A group of approximaiely 30 individuals occupied a sheltered hollow between two 
moss-covered buttress roots of an unidentified canopy tree along a ridgetop trail (Fig. 2). 
The roost was found again in the same location the following year (July and August 
1984), when a preliminary investigation of moth behavior at the roost was undertaken. 
No other Ocalaria roosts were observed in either 1983 or 1984, but in 1985 several roosts 
were found in the immediate area (including the original site), as well as on other ridges. 

The number of moths found at the roost in 1984 varied, but on average 18-21 indi- 
viduals occupied the roost during the day, as estimated by visual censuses made during 
10 observation periods. The moths typically congregated during the day in the hollow 
portion between the buttress roots in a loosely clustered group about 1 m above ground 
(Fig. 3). While roosting the moths were motionless, held their wings in a partially upright 
position, and were well camouflaged against the textured, mossy trunk in the shady 
hollow (Fig. 4). Most individuals would not move unless prodded. Once when attempting 
to capture a specimen, we disturbed two of the moths. They flew out of the hollow and 
landed on the same tree about 60° around the trunk. One individual was immediately 
seized and eaten by an Anolis sp. lizard (Anolidae), predation similar to that described 
for the diurnally roosting skipper, Celaenorrhinus fritzgaertneri (Hesperiidae) (DeVries 
et al., cited above). 

To document moth activity at the roost, and times when the roost was occupied and 
not occupied, two evening and two morning censuses were undertaken. A synchronous 
pattern similar to that observed in the cave-roosting skipper C. fritzgaertneri character- 
ized both departure and return to the roost. In July and August at Parque Nacional 
Corcovado the sun sets at approximately 1745 h, and by 1810 h it is completely dark. 
On August 12, observations were begun at 1730 h. Twenty-two moths were found in the 
roost at that time. As nightfall approached, the moths appeared restless, some changed 


VOLUME 40, NUMBER 2 125 


tok 





= £3 
eke t 
% 
gf, te 
Po. 


“ot ae 9 sue alte 


Fics. 1-4. 1, Dorsal surface (top) and ventral surface (bottom) of a male Ocalaria 
undescribed species from Costa Rica; 2, Buttressed tree where Ocalaria roost was ob- 
served; 3, A cluster of Ocalaria moths in the roost; 4, Detail of roosting Ocalaria indi- 
vidual. 


position in the roost. At 1750 h two moths left their perch and began to flutter inside the 
hollow. A few seconds later these moths flew out of the roost. Shortly thereafter (1755 
h) most of the roosting moths followed suit, first hovering from 5 to 15 seconds in the 
hollow before flying out into the night. By 1800 h all had left the roost. At this time the 
light was too dim to read handwritten notes. The departing moths, which have a weak, 
fluttery flight, were not observed to land on the adjacent vegetation, and these moths 
were never found in collections made at lights in 1984. It is not known how far from 
the roost they fly or what they do during the night, but they did not return to the roost 
until daybreak. A second observation period on August 15 showed the same pattern. 
On the mornings of August 14 and 18, the moths’ return to the roost was observed. 
Observations began at 0430 h when it was still completely dark, and no moths were 
found on the roost on either occasion. The first light of dawn began at about 0500 h, and 
thereafter light intensity increased rapidly until about 0535 h when the sun rose above 
the horizon; however, light levels remained low in the forest understory where the roost 
was located until nearly 0600 h. On August 14 the first moth to approach the roost came 
at 0512 h and landed almost immediately. At this time light levels were barely high enough 
to distinguish handwriting on a page. Three more moths entered the hollow at 0515 h, 
and spent 10-30 seconds hovering in the hollow before landing on the trunk. As the sky 
became lighter (0525 h) a large pulse of moths arrived at the roost, with several fluttering 


126 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


simultaneously in the hollow for 5-20 seconds before taking positions on the trunk. By 
0545 h all activity ceased, and the 18 individuals in the roost remained stationary. No 
more moths were seen to arrive from that time until 0615 h when the observation period 
ended. A similar pattern of arrival was observed on the morning of August 18. 

These observations suggest that the timing of roost departure and arrival in Ocalaria 
may be closely tied to light intensity, as found in cave-roosting skippers (DeVries et al., 
cited above). Our observations suggest that roosting sites are important for Ocalaria since 
moths were found at the same unremarkable roost site for two consecutive years, with a 
fairly stable roost membership over the four-week observation period during both years. 
This is corroborated by further observations in Costa Rica and Panama during 1985-86, 
where the same roosts were found occupied by populations of Ocalaria over long periods. 
Recovery of marked individuals from the roost where they were originally captured 
(mark-release-recapture experiments conducted in 1985) indicates that Ocalaria, like the 
cave-roosting skippers, are “faithful” to a particular roost (Greig & DeVries, in prep.). 

Although there are parallels in the roosting behavior between this species and the 
skipper butterfly Celaenorrhinus fritzgaertneri, there is little evidence that Ocalaria 
moths are roosting as an aestivation response. The roosts were first observed during the 
early wet season when larval resources are presumably plentiful (however, there are no 
host plant data available for any species in the genus), and our subsequent observations 
in both Costa Rica and Panama suggest that Ocalaria roosts may persist throughout the 
year. 

We thank the Servicio de Parques Nacionales de Costa Rica and the personnel of 
Parque Nacional Corcovado for hospitality during our stay in Costa Rica. J. Rawlins 
identified the moth to genus and R. Poole confirmed that it is an undescribed species. I. 
F. B. Common and an anonymous reviewer provided helpful comments on the manu- 
script. 


NANCY GREIG, Division of Biological Sciences, AND P. J. DEVRIES, Department of 
Zoology, University of Texas, Austin, Texas 78712. 


ANNOUNCEMENT 


SUPPLEMENTAL KEY WORDS FOR JOURNAL PAPERS 


Starting with Journal volume 41 (1987), up to five index or key words or phrases not 
already in titles will appear with articles and general notes. Such words should accom- 
pany manuscripts being submitted now. Terms should be selected that, together with the 
title, most fully describe a paper’s scope and depth. Authors are best able to select 
supplemental index words, but reviewers and editors can help. A comprehensive index- 
word resource is R. H. Foote’s 188-page Thesaurus of Entomology, published by the 
Entomological Society of America in 1977. 

Supplemental key words will aid the skimming reader. But more importantly, they 
will be used to augment and enrich the index appearing at the end of each Journal 
volume. Recent indexes have been limited to little more than species, genus, family, and 
author. The wealth of Journal content pertaining to topics such as geographic variation, 
host relations, phenology, predation, and reproduction, has gone largely unindexed. 

Supplemental key words will make more kinds of information retrievable, and result 
in greater Journal usefulness. 


WILLIAM E. MILLER, Editor 


Journal of the Lepidopterists’ Society 
40(2), 1986, 127 


CLARIFICATION OF THE LARVAL HOST PLANT OF 
EPIDEMIA MARIPOSA (LYCAENIDAE) IN NORTHERN CALIFORNIA 


The larval host plant of Epidemia mariposa (Reakirt) has long been thought to be 
Polygonum douglasii Green (Polygonaceae) (Tietz 1972, An index to the described life 
histories, early stages, and hosts of the Macrolepidoptera of the continental United States 
and Canada, A. C. Allyn, Sarasota, Florida, 426 pp.; Pyle 1981, The Audubon Society 
field guide to North American butterflies, Alfred A. Knopf, New York, 916 pp.). Other 
Epidemia, especially the closely related E. nivalis (Boisduval), use Polygonum douglasii 
as host plants (Pyle, cited above; Howe 1975, The butterflies of North America, Double- 
day and Co., Garden City, New York, 633 pp.). However, other studies suggest that 
Vaccinium (Ericaceae) L. is the host plant of E. mariposa. In a paper describing the egg 
of E. mariposa (Coolidge 1910, Can. Entomol. 42:316), its host plant was recorded as 
Vaccinium in Yosemite. Although this record probably was for E. mariposa, the paper 
referenced by Coolidge (Lembert 1894, Can. Entomol. 26:46) recorded the butterfly that 
oviposited on Vaccinium as Chrysophanus arota (Boisduval) which uses Ribes or Goose- 
berry. Our recent field and laboratory studies confirm that Vaccinium is the larval host 
plant for E. mariposa in northern California. 

On 4 August 1984 we visited Cedar Lake (1,700 m elev., 24 km southwest of Mt. 
Shasta), Siskiyou Co., California, to obtain females of E. mariposa for life history studies. 
No Polygonum douglasii was observed, yet P. bistortoides Pursh. was found in and 
around the periphery of the bog, and in neighboring marshy areas. A number of potential 
host plants including P. bistortoides were taken from the area for ovipositional studies. 

Eight females of E. mariposa were put in oviposition chambers with varying combi- 
nations of plants collected from the bog, along with either Rumex californica Rech. or 
R. crispus L. (Polygonaceae). One female was exposed to both Rumex crispus and Vac- 
cinium arbuscula (Gray) Merriam. Only two eggs were oviposited, both on V. arbuscula. 
Four of the eight females were switched to V. arbuscula, while the remaining females 
were left on the Rumex and Polygonum. The following day, more than 50 eggs were 
oviposited by the females switched to V. arbuscula, while none were oviposited on the 
Rumex or Polygonum. 

After being left at room temperature for a month, the eggs were refrigerated at 4°C. 
They were removed after two months, and allowed to warm to room temperature (late 
November 1984). Within two weeks, 10 ova hatched and the neonatal larvae were con- 
fined with both Rumex crispus and Vaccinium corymbosum L. (nursery stock). Larvae 
refused to feed on R. crispus, which we have found acceptable to E. nivalis, yet fed on 
fresh young shoots of V. corymbosum. Eggs sent to John F. Emmel were reared to 
maturity on Vaccinium corymbosum. 

On 19 June 1985, we returned to Cedar Lake to look for larvae of E. mariposa on 
Vaccinium. By beating branches into a heavy cloth net, we obtained two mature larvae 
from Vaccinium arbuscula, but none from V. occidentale Gray, which also occurs in the 
bog. Perhaps E. mariposa females oviposit on particular species of Vaccinium; the but- 
terflies are not found in all areas where Vaccinium is abundant within their range. 

A fresh female E. mariposa collected 27 July 1985 at Tioga Pass, Mono Co., California, 
was induced to oviposit on the local Vaccinium, V. nivictum Camp. Vaccinium nivictum 
is considered to be a close relative of V. arbuscula (at one time both species were 
considered the same as V. caespitosum Michx.). Unfortunately, the eggs were infertile. 

Voucher specimens of E. mariposa from Cedar Lake are in the authors’ collections 
and the collection of the Entomology Department, University of California at Riverside. 
Specimens of the Vaccinium are in the University’s herbarium. 


GORDON F. PRATT AND GREGORY R. BALLMER, Department of Entomology, Univer- 
sity of California, Riverside, California 92521. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 128-129 


POLYCHRYSIA MORIGERA (NOCTUIDAE) TRAPPED IN A SOUTHERN 
LADY’S-SLIPPER LABELLUM IN TENNESSEE 


Catling (1974, Newsletter Mich. Entomol. Soc. 19(1):1, 3) conjectured that a skipper 
butterfly, Thymelicus lineola, might exemplify a case in which a species recently intro- 
duced into North America had not “learned” that entering labella of the showy (or 
queen) lady’s-slipper flowers was both unproductive of nectar and likely to result in 
entrapment. The discovery described below extends his hypothesis to the southern lady’s- 
slipper, Cypripedium kentuckiense Reed, and a plusiine noctuid moth, Polychrysia mor- 
igera (Edwards). The moth is a western American species which seems to have recently 
become established in the East. 

On 24 May 1984, Medley discovered two moths inside the shoelike labellum of one 
flower of the southern lady’s-slipper in Scott Co., Tennessee. The plant was located at 
the margin of a mesic floodplain forest, and was growing in a more open area than were 
most of the others in that population. One moth was dead, the other still alive; both were 
slightly worn, probably from their efforts to escape. This was the first discovery of a 
lepidopteran in flowers of this species during a study of lady’s-slippers populations in 
which over 1,000 blossoms were examined in Kentucky, Tennessee, Arkansas, and Okla- 
homa from 1980-85. That number includes about 200 at the Scott Co. site examined at 
the time the moths were recovered. The only other insect found in the overall survey 
was a single large bee (Xylocopa sp.?) in a labellum in Jefferson Co., Arkansas. 

Covell identified the moths as male Polychrysia morigera, a species previously restrict- 
ed to far western states (Eichlin & Cunningham 1978, U.S. Dep. Agr. Tech. Bull. 1567, 
122 pp.). However, it appeared in Kentucky in 1976 (Covell determination), and is now 
known from five Kentucky counties, collected from 20 May to 13 June. The only other 
known eastern record is a recent capture in Pennsylvania (J. G. Franclemont, pers. 
comm.). It is unknown from Missouri, which has been extensively surveyed for Lepidop- 
tera (J. R. Heitzman, pers. comm.). The present record constitutes the first report of 
moths in southern lady’s-slipper flowers, and also the first Tennessee record of P. mori- 
gera. 

Trapping of Lepidoptera by flowers of other Cypripedium species has previously been 
reported (Arthur 1962, Proc. Entomol. Soc. Ont. 92:190-191; Stoutamire 1967, Mich. 
Bot. 6:159-175; Catling, cited above; Barrows 1983, J. Lepid. Soc. 37:265-267). Small 
bees rather than Lepidoptera appear to be pollinators of Cypripedium species (Stouta- 
mire, cited above). Catling explained how no nectar is produced by the flowers, but odor 
and color attract bees into the lip. They then exit through small openings placed so as to 
force the bees to rub against pistil and anthers. These openings are too small to permit 
escape by butterflies. Based on a literature review, Stoutamire summarized insect rela- 
tions with five Cypripedium species. None involved C. kentuckiense. The only moth 
record was for Tetracis cachexiata Guenée (Geometridae; published as T. lorata Grote, 
a synonym), reported by J. Newman to be resting on the labellum of a stemless lady’s- 
slipper, Cypripedium acaule Ait., in Michigan. The only butterflies observed inside la- 
bella of any species were recorded in those of showy lady’s-slipper, C. reginae (Walt.). 
These were the following skippers (Hesperiidae): Thymelicus lineola (Ochsenheimer), 
Epargyreus clarus (Cramer), Polites mystic (Edwards), and Polites themistocles (La- 
treille). Arthur (cited above) found six individuals of the European skipper, Thymelicus 
lineola, in labella of C. reginae in Ontario. Catling found the same skipper in “at least 
half” of the “about 100” flowers of showy lady’s-slipper that he examined in July 1971, 
in Grey and Simcoe counties, Ontario. Some blossoms contained as many as five butterflies 
each, some alive, some dead. In early June 1972, Catling and colleagues found one flower 
containing a European skipper in Washtenaw Co., Michigan. Catling noted that other 
skipper species of similar size were more abundant in these habitats than T. lineola, but 
none found in flowers. Barrows (cited above) found up to 24 T. lineola in a single C. 
reginae labellum in Cheboygan Co., Michigan. He recovered 646 T. lineola, as well as 
other insects, including “geometrid moths” from an undisclosed number of blossoms 
examined. Thymelicus lineola and unidentified geometrids were also found in C. cal- 


VOLUME 40, NUMBER 2 129 


ceolus L., which we determined from a Barrows illustration to be the taxon now consid- 
ered Cypripedium parviflorum Salisb. Barrow’s 1978 samples consisted of 73-90% males. 

Catling conjectured that because T. lineola had recently been introduced into North 
America from Europe (1910), it might not have undergone microevolution enabling it 
to avoid entering and becoming fatally trapped in lady’s-slipper flowers. The hypothesis 
is plausible in light of the many trapped T. lineola he and Barrows reported, contrasting 
with a lack of common “native” skippers trapped in labella. In support of Catling’s 
hypothesis, we suggest that P. morigera may also have become established in eastern 
North America recently, and might also be unfamiliar with the dangers of entering these 
flowers—a case parallel to that of the European skipper farther north. 

We thank J. G. Franclemont, Cornell University, and J. R. Heitzman, Independence, 
Mo., for their input; and W. Herb Wagner, University of Michigan, for reviewing this 
manuscript. 


CHARLES V. COVELL, JR. AND MAX E. MEDLEY, Department of Biology, University 
of Louisville, Louisville, Kentucky 40292. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 129 


AVIAN PREDATION OF ALPINE BUTTERFLIES 


Because direct observation of predation on butterflies is exceedingly rare (Bowers et 
al. 1985, Evolution 39:93-103), the following observations of avian predation on a variety 
of alpine butterflies may be of interest. 

While studying the foraging ecology of nesting water pipits (Anthus spinoletta (L.)) 
on the Beartooth Plateau (elev. 3,300 m), Park Co., Wyoming, I observed a nesting female 
pipit capture and consume the following butterflies from 6-10 August 1983, all between 
1721 and 1985 MDT: two Speyeria mormonia (Boisduval), one Parnassius phoebus 
Fabricius, and one Euphydryas editha (Boisduval). In each case the butterfly was flushed 
from the ground as the pipit foraged by walking through the tundra vegetation. The 
wings were torn or flicked off before the body was eaten. 

That one bird was seen capturing and eating four butterflies in a relatively short time 
(10 h of observation) suggests that avian predation could at times cause important mor- 
tality in some alpine butterfly populations. The proposed impact of avian predation on 
alpine butterflies would probably not be as extreme as that documented for other, low 
elevation, butterflies (Calvert et al. 1979, Science 204:847-851; Fink et al. 1983, Biotro- 
pica 15:151-158; Bowers et al. 1985, Evolution 39:93-103). However, it might still be a 
significant factor in the demographics of narrowly distributed taxa, such as Boloria ac- 
rocnema Gall & Sperling (Gall 1984, Biol. Conserv. 28:111-138), especially if the breed- 
ing density of alpine birds is high where the butterflies are concentrated. 

My field work was funded by Sigma Xi, a Bertha Morton Scholarship, and the De- 
partment of Zoology, University of Montana. 


PAUL HENDRICKS, Department of Zoology, University of Montana, Missoula, Mon- 
tana 59812. Present address: Department of Zoology, Washington State University, 
Pullman, Washington 99164-4220. 


Journal of the Lepidopterists’ Society 
40(2), 1986, 130 


OVIPOSITIONAL RESPONSE OF EUREMA NICIPPE (PIERIDAE) TO 
PHYLLANTHUS TENELLUS (EUPHORBIACEAE) 


The host plants for Eurema spp. are mostly members of the legume subfamilies Mi- 
mosoidae and Caesalpinoidae. Most of the other members of the pierid subfamily Co- 
liadinae feed on other plants in the legume family (Fabaceae) (Howe 1975, The butter- 
flies of North America, Doubleday, New York). All records associated with Eurema larvae 
in the collection at the University of Florida are for various species of Cassia (Caesal- 
pinoidae) (Habeck, unpubl. data). Thus I was surprised to observe adult female Eurema 
nicippe (Cramer) alight in a dense patch of the exotic Euphorbiaceae Phyllanthus te- 
nellus Roxb. and display what appeared to be ovipositional behavior. During a three- 
hour period (0930-1230 h) on 7 October 1985 near Gainesville, Florida, I observed 46 
instances of this behavior; I expected to find eggs on the Phyllanthus stems visited by 
the butterflies, and actually found 6 eggs. These were kept in petri dishes with fresh 
Phyllanthus; hatching larvae did not feed and wandered around the dishes. When offered 
leaves of Cassia chamaecrista L. (partridge-pea) three days later they immediately began 
feeding. 

The leaves and growth form of Phyllanthus tenellus, a frequent weed of moist, shaded 
disturbed ground, strikingly resemble a usual host plant in northern Florida, Cassia 
chamaecrista (pers. observ. and Kimball 1965, The Lepidoptera of Florida, Division of 
Plant Industry, Florida). Cassia chamaecrista has pinately compound leaves, while the 
small leaves of Phyllanthus tenellus are closely arranged in a single plane along elongate 
side branchlets. This suggests that E. nicippe locates its host plant first by sight, but that 
because oviposition is infrequent on this incorrect host plant, secondary chemotactic 
responses are involved. 

Another common host of Eurema spp. (including E. nicippe) in Florida is Aeschy- 
nomene americana L. (joint vetch), also a legume, but a distant relative of Cassia. Few 
of the other legume species are used by Eurema spp. The morphology of A. americana 
also strongly resembles that of Cassia chamaecrista. I propose that Aeschynomene has 
become a regular host plant of Eurema because of continued ovipositional mistakes on 
this common plant, and the taxonomic relatedness of Aeschynomene and Cassia. Could 
Phyllanthus eventually become a host plant for Eurema nicippe? If taxonomic related- 
ness is necessary for host switching to occur, probably not. But if biochemical similarity 
or an altered detoxifying system are possible, then host switching across wider taxonomic 
barriers might occur. If not, an improved host recognition in Eurema butterflies should 
evolve. 

Within the genus Cassia there is a great variety of leaf morphologies. This diversity 
in leaf morphology could have evolved in response to the several genera and many species 
of Cassia-feeding pierids that rely on visual clues for host finding. 


MICHAEL J. PLAGENS, 4407 E. Lee, Tucson, Arizona 85712. 


Date of Issue (Vol. 40, No. 2): 23 October 1986 


EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
BoyYcE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Short manuscripts concerning new state records, current events, and notices 
should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence, KS 66044 
U.S.A. Journal contributors should prepare manuscripts according to.the following in- 
structions. 

Abstract: An informative abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name with author. Mea- 
surements should be given in metric units; times should be given in terms of the 24- 
hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. References 
to footnotes should be numbered consecutively, and the footnotes typed on a separate 
sheet. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In general notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30). 

Illustrations: Only half of symmetrical objects such as adults with wings spread 
should be illustrated, unless whole illustration is crucial. Photographs and drawings should 
be mounted on stiff, white backing, arranged in the desired format, allowing (with 
particular regard to lettering) for reduction to their final width (usually 4% inches). 
Illustrations larger than 8% x 11 inches are not acceptable and should be reduced pho- 
tographically to that size or smaller. The author’s name, figure numbers as cited in the 
text, and an indication of the article’s title should be printed on the back of each mounted 
plate. Figures, both line drawings and halftones (photographs), should be numbered 
consecutively in Arabic numerals. The term “plate” should not be employed. Figure 
legends must be typewritten, double-spaced, on a separate sheet (not attached to the 
illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to 
each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the 
proofs. A page charge of $60 per full Journal page is applicable to all authors. Unfunded 
authors may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 





PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


Wuy PIERIS RAPAE IS A BETTER NAME THAN ARTOGEIA RAPAE 
(PIERIDAE). Robert K. Robbins & Pamela M. Henson ........ 


RESTINGA BUTTERFLIES: BIOLOGY OF SYNARGIS BRENNUS (STICH- 
EL) (RIODINIDAE). Curtis J. Callaghan 2... oe 


MALE AND FEMALE GENITALIA OF PHOEBIS EDITHA (BUTLER): 
How THEY DIFFER FROM HISPANIOLAN P. SENNAE 
(LINNAEUS) (PIERIDAE). John G. Coutsis 2a 


GENUS DIPTYCHOPHORA ZELLER AND A RELATED NEW GENUS 
STENEROMENE FROM THE NEOTROPICAL REGION (PYRALIDAE: 
CRAMBINAE). David E. Gaskin 


GENERAL NOTES 


Small-Nicolay collection to Smithsonian. Robert K. Robbins & J. F. Gates 
Clarke: Vs) ole Vie Scorpion i tdercnnnteelarcaricoerntvpaet le an ne 


Observations on the diurnal gregarious roosting of Ocalaria sp. (Noctuidae) 
in Costa Rica. Nancy Greig & P: J. DeVries |... ae 


Clarification of the larval host plant of Epidemia mariposa (Lycaenidae) in 
northern California. Gordon F. Pratt & Gregory R. Ballmer .............. 


Polychrysia morigera (Noctuidae) trapped in a southern lady’s-slipper label- 
lum in Tennessee. Charles V. Covell Jr. t¢ Max E. Medley ................. 


Avian predation of alpine butterflies. Paul Hemdbricks 22... cctecsssssecssssssenseeeeeeeoes 


Ovipositional response of Eurema nicippe (Pieridae) to Phyllanthus tenellus 
(Euphorbiaceae). Michael J. Plagens 2.002 


BOOK REVIEW. Se ee m4 


ANNOUNCEMENT 
Supplemental key words for Jotrmal papers ooo. iceicscesseccsseseceeeseeneeeseesncessesnseeceesnsnesensanee 


130 
123 


Volume 40 1986 Number 3 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





Ss o 


} x Nae > AF 
~ > BY + 12 we pits 2, 7 
AD git ale ashes eS 


23 January 1987 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


DoucLas C. FERGUSON, President GERARDO LAMAS M.., Vice President 
CLIFFORD D. FERRIS, Immediate Past President EBBE SCHMIDT NIELSEN, Vice Presi- 
OLAF H. H. MIELKE, Vice President dent 

RICHARD A. ARNOLD, Secretary Eric H. METZLER, Treasurer 


Members at large: 


BoycE A. DRUMMOND III MIRNA M. CASAGRANDE M. DEANE BOWERS 
JOHN LANE EDWARD C. KNUDSON RICHARD L. BROWN 
ROBERT K. ROBBINS FREDERICK W. STEHR PAUL A. OPLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “‘to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229-1306, U.S.A.; and address changes to: 
Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 
90007-4057 U.S.A. 


To obtain: 

Back issues of the Journal and NEWS (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society’) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, CA 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Boulevard, 
Los Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Soci- 
ety, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 


Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from 
Dyar 1899, J. New York Entomol. Soc. 7:61-67. Suggested by Marc E. Epstein. 


JOURNAL OF 


Tue LeEeriporrtreERISTS’ SOCIETY 
Volume 40 1986 Number 3 


Journal of the Lepidopterists’ Society 
40(&), 1986, 1381-137 


A SIMPLE INSTANT DIET FOR REARING 
ARCTIIDAE AND OTHER MOTHS 


RAINER BERGOMAZ AND MICHAEL BOPPRE! 


Universitat Regensburg, Zoologie - SFB4/B6, UniversitatsstraBe 31, 
D-8400 Regensburg, Federal Republic of Germany 


ABSTRACT. A bean-based semiartificial diet was developed empirically for rearing 
species of Rhodogastria and Creatonotos (Arctiidae), foodplants for which were un- 
available or unknown. It was also used successfully for Heterocera in 27 genera of five 
families, and is likely suitable for rearing additional species of moths. Preparation of the 
diet is simple even in the field, and its components are inexpensive. 


There are many practical reasons for attempting to rear insects on 
artificial diets, in particular to provide food independent of place or 
season. Numerous diet recipes have been published (Vanderzant 1974, 
Singh 1977, Singh & Moore 1985), most of which are based on either 
wheatgerm, wheatgerm + casein or beans. Their diversity arises main- 
ly from their various sources of vitamins, proteins, carbohydrates, fatty 
acids, and preservatives. Most published diets for Lepidoptera have 
been tested with only one or a few closely related species, and rearing 
success can seldom be standardized to permit fully objective compari- 
sons. The suitability of a certain diet for any given but untried species 
is thus not predictable. 

Faced with the need to breed Rhodogastria and Creatonotos (Arc- 
tiidae), we considered artificial diets because the natural foodplants 
were either unknown or not continuously available. We also wanted a 
simple medium for use under field conditions in the tropics. To these 
ends, we modified recipes from the literature in a largely empirical 
fashion until we ended with an instant diet, which is cheap and easily 
prepared even during field work in the tropics. This simple diet also 


‘Present address of authors: Forstzoologisches Institut der Universitat Freiburg, Fohrenbiih] 27, D-7801 Stegen- 
Wittental, West Germany. 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Recipe of semiartificial diet. For simple preparation, 6 parts of I, 1 part of 
agar, and 20 parts of water are combined. 


Amount 

Ingredients %} g 
Bean flour® 15.0 75.0 
Brewer's yeast 3.5 ee 
Ascorbic acid 0.7 3.5 
Cholesterol 0.1 0.5 I 
Sorbic acid 0.1 0.5 
Methyl-p-hydroxybenzoate 0.1 0.5 
Streptomycin 0.08 0.4 
p-formaldehyde — 0.15 
Formaldehyde (10%) 0.3 
Germ oil with a-tocopherol! 0.7 
Agar agar® 3.0 15.0 
Water 76.0 381.0 


! Weight/weight, except formaldehyde, which is volume/volume. 

2 To yield ca. 500 g of diet. 

3 Dried white beans (“WeiBe Bohnen,’ Phaseolus compressus var. albus) obtained from a supermarket and pulverized 
by coffee grinder. 

4 10 g a-Tocopherol + 140 ml germ oil (“Mazola® Keimdl,” a salad oil produced and distributed by Maizena GmbH, 
Heilbronn). 

5 High gel-strength powder, research grade (SERVA, Heidelberg, New York). 


appeared well suited to rearing other arctiids and a variety of species 
of other heteroceran families. 


MATERIALS AND METHODS 
Diets 


Diets initially tested were based on wheatgerm, and were similar to 
those described by Vanderzant (1967) and Bell and Joachim (1976). 
These we modified by using frozen, tinned, or dried beans (Vicia faba, 
Soja hispida), testing mixtures of beans and wheatgerm in different 
proportions, and adding proteins, salts, and vitamins as well as leaf- 
powders in varying amounts. Eventually, we omitted several standard 
ingredients of artificial diets to create a simpler recipe. The diet we 
found best (Table 1) is prepared by parboiling water, and adding the 
agar while stirring. Other ingredients are then mixed thoroughly (pref- 
erably with an electric hand mixer) with the gelling agar. Flat plastic 
dishes are filled with this pap to a depth of 4 cm. Cooling and drying 
is allowed for up to 12 h at room temperature before storage in a 
refrigerator. With refrigeration, this diet can be stored for at least five 
weeks; in the field, smaller amounts of diet were prepared to last for 
three to five days. 

After obtaining good rearing results with this diet, we modified it 
slightly to produce an instant diet which could be prepared more eas- 
ily, particularly during field work in the tropics. The only departure 
from the original recipe is that 0.08% p-formaldehyde (solid) was sub- 


VOLUME 40, NUMBER 3 ies 


stituted for formaldehyde solution; also, in the field, germ oil and 
a-tocopherol were omitted. Agar and a mixture of all other solid in- 
gredients were put separately into two plastic containers, which allows 
for storage and transport (mailing) without refrigeration. When the diet 
was required, it was prepared by merely mixing agar with boiling water 
and adding the mixture (and oil) while stirring thoroughly. In practice, 
we use a 10 ml! plastic container to measure 1 part of agar, 6 parts of 
the mixture, and 20 parts of water (or multiples thereof if more diet is 
required). However, these proportions work only if the beans are ground 
finely (as coffee for filtering), and high gel-strength agar (SERVA, 
Heidelberg, New York) is used; otherwise, different weight-to-volume 
relationships have to be determined. 


Insects 


Rhodogastria phaedra (Weymer) from Kenya (East Africa) and 
Creatonotos transiens (Walker) from Sumatra, Indonesia, were exten- 
sively investigated. Stocks originated from inseminated females sent to 
our laboratory. The foodplants of Rhodogastria phaedra are unknown, 
and the few known for other Rhodogastria are not available to us. 
Creatonotos transiens is a polyphagous species which we had previ- 
ously reared successfully on European substitute foodplants such as 
Taraxacum officinale L. 

The suitability of the simplest recipe (the instant diet) was tested 
with several European and tropical Lepidoptera (Table 2) to which we 
had unexpected access. 

In routine cultures, as many as 50 neonate larvae were put into a 
clear plastic Petri dish (94 mm X 16 mm) containing a piece of diet 
(about 6 X 6 X 3 mm). Moistened filter paper was sometimes added. 
Later instars were kept in groups of 10 to 380 in plastic containers 
(200 x 200 x 95 mm), and given larger (but always 5 mm thick) pieces 
of diet. Every third day, larvae were transferred to a clean container 
with fresh diet. For quantitative rearing data, larvae were sometimes 
kept singly in Petri dishes. 

We usually assessed the success of a diet by comparing the visual 
appearance of adults with field-caught specimens, and by checking the 
fecundity of females and the fertility of their eggs. In some cultures 
we also compared diets by weighing pupae, counting eggs, and mea- 
suring rates of development. For reasons given below, however, we 
report only on our final results, and omit the presentation of detailed 
data. 


RESULTS 


The diets of Vanderzant (1967) and Bell and Joachim (1976) were 
only moderately successful with Rhodogastria and Creatonotos. Better 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


success was achieved by using beans instead of wheatgerm, while sub- 
stituting casein by albumin and/or eggs as a protein source did not 
have an obvious effect. Using Phaseolus beans as a base, we stepwise 
left out casein, Wesson’s salt (compare Singh 1977) and/or vitamin 
solutions without reduced rearing success. Later, sucrose, choline chlo- 
ride, and inositol were found to be unimportant additions. The resulting 
best recipe from our trials is given in Table 1. 

With Creatonotos in particular, we noted a dramatic group effect: 
larvae kept individually resulted in pupae which were up to 30% light- 
er than those of larvae kept in groups of 15-20. Thus, optimal rearing 
does not always depend on diet composition alone; indeed, other breed- 
ing conditions can be of overriding importance. 

Notable general observations were, briefly: 1. White beans (Phase- 
olus compressus var. albus) appeared to be a better base for artificial 
diet than horse-beans (Vicia faba); this finding supports the experience 
of El-Guindy et al. (1979). 2. Soaking beans (previously done by most 
authors, such as Shorey & Hale 1965) proved unnecessary; pulverizing 
the seeds greatly simplifies preparation. 3. The use of methyl-p-hy- 
droxybenzoate and sorbic acid enables extended storage of diet without 
refrigeration, and combats insect pathogens. Their concentration should 
nevertheless be kept as low as possible, and they can be omitted if the 
culture is healthy, and the diet is replaced frequently. 4. The quanti- 
tative composition of diet ingredients, particularly of water, appears 
less important than we originally judged from the literature. 

Larvae of 38 species of moths, belonging to 27 genera of five families 
(Table 2) accepted our simple diet and developed with no unexpected 
mortality to adults that did not differ noticeably in size, color, and 
other characteristics from field-caught material. Females invariably 
proved fecund and eggs fertile. Rearing success from newly hatched 
larvae to adults was 80-100% in all species. Most species were reared 
for at least four generations (C. transiens for 20) without exhibiting 
apparent changes. However, larvae of sevaral species, including Eu- 
chromia amoena (Moschler) (Ctenuchiidae), Antherea pernyi Buerin- 
Meneville (Saturniidae), and all Rhopalocera, refused the diet. When 
checking the suitability of the simple diet with the species listed above, 
we experienced the following: 1. Preservatives may function as 
antifeedants; in some species we observed refusal if the concentration 
of methyl-p-hydroxybenzoate was higher than 0.1%. Hirai (1976) ob- 
tained similar results, and also reported noxious effects of sorbic acid. 
2. When nonbacterial pathogens affected our cultures, we cured the 
larvae by adding 0.15% Fumidil B (Abbot Laboratories), a pharma- 
ceutical marketed by CEVA, Paris, to treat honeybees for Nosema 
infection. 3. Using diets can cause a delay of two to three days before 


VOLUME 40, NUMBER 3 135 


TABLE 2. Species successfully reared on the simple diet. 


Species Origin 
Arctiidae 
“Amsacta”’ emittens Walker Sri Lanka 
Arctia caja L. Germany 
Ammobiota festiva Hufnagel Turkey** 
Callimorpha principalis Coll. China** 


Creatonotos gangis (L.) Sumatra, India 
Creatonotos transiens (Walker) Sumatra, Java 
Cycnia mendica Cl. Germany** 


Diacrisia sannio L. Germany 
Nyctemera sp. India* 
Ocnogyna joiceyi Talbot Morocco** 
Orodemnias cervini Fallou Germany** 
Paralacydes sp. Kenya 
Pericallia ricini Fabr. India* 
Phragmatobia fuliginosa L. Germany 
Rhodogastria bubo (Walker) Kenya 
Rhodogastria carneola Hampson Kenya 
Rhodogastria luteibarba Hampson Kenya 
Rhodogastria phaedra (Weymer) Kenya 
Rhodogastria thermochroa Hampson Kenya 
Rhodogastria vitrea (Plétz) Kenya 
Rhodogastria n. spp. (4) Kenya 
Spilosoma sp. Borneo 
Spilosoma menthastri Esp. Germany 
Teracotona rhodophaea Walker Kenya 
Utetheisa pulchella L. India* 
Ctenuchiidae 
Pseudonacia sp. Kenya 
Sphingidae 
Manduca sexta L. America 
Noctuidae 
Achaea lienardi Boisd. Kenya 
Autographa gamma L. Germany 
Paradiarsia glareosa Esp. Germany** 
Polymixis serpentina Tr. Yugoslavia** 
Staurophora celsia L. Germany** 
Lymantriidae 
Euproctis lunata Hb. India* 
Olene (Dasychira) mendosa Hb. India* 
Pralis securis Hb. India* 


* M. Eckrich, pers. comm. 
** K. Heuberger, pers. comm. 


newly hatched larvae begin feeding. 4. Diet is best accepted by neonate 
larvae, and it is often difficult or impossible to switch from foodplant 
to diet. 5. Addition of leaf powder to the diet (such as Taraxacum 
officinale for A. gamma) caused some neonate larvae to start feeding 
earlier than on diet lacking plant material, but always increased the 


136 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mortality of the larvae, and greatly prolonged development. 6. Color 
might be a cue in food detection and acceptance. Larvae of Autogra- 
pha in a choice experiment preferred diet colored green by chromoxide 
(compare McGinnis & Kasting 1964). However, this did not result in 
better rearing success. 


DISCUSSION 


We succeeded in our primary objective of rearing certain tropical 
moths whose foodplants are unknown or unavailable, and we found a 
diet suited to a wide spectrum of species. The diet we report not only 
enabled us to discover the features of previously unknown larvae, so 
as to facilitate search for their natural foodplants, but also to obtain 
adults for breeding and experimental studies. 

Existing reports of rearing arctiids on artificial diets include those of 
Gottel and Philogene (1978) for Pyrrharctia isabella (J. E. Smith), Vail 
et al. (1967) for Estigmene acrea (Drury), Singh (1977) for Hyphantria 
sp., Conner et al. (1981) for Utetheisa ornatrix, Bathon (1977) for 
Spilosoma maculata Stoll., and Moreau (1965) for Arctia caja L. Most 
of these diets are wheat-based. In contrast, our experience with Arc- 
tiidae showed that bean-based diets yield superior results. Apparently, 
beans have higher nutritive value for lepidopterous larvae than wheat- 
germ. Bean-based diets may also be easier to prepare, and are less 
expensive. Salts, vitamins, certain proteins, and other ingredients used 
by other authors can be omitted from bean-based diets, reducing costs 
significantly (1 kg of cur instant diet costs ca. 7 Deutsch-Marks). In 
broad comparisons among reports, it must nevertheless be understood 
that significant differences occur in the quantitave and qualitative 
composition of nutrients between different species/strains of beans 
(Schlieper 1982). 

Our medium turned out to be similar to the one described for noc- 
tuids by Shorey and Hale (1965). We have not compared our results 
with those of others because not only do the recipes differ, but also the 
species and rearing conditions—variables often as important as diet 
composition. Furthermore, due to the lack of natural foodplants for 
most species, we could not compare cultures on plants with those on 
diets. 

We cannot explain why our diet proved successful for the variety 
of species tested. Though our results are certainly open to further re- 
finement, we here report our findings as they stand now, and briefly 
discuss our experience because our instant diet is suited to a variety of 
unrelated species with different foodplant requirements. Also, in con- 
trast to many other diets, it is easy to prepare, and inexpensive, and 
thus may help other workers, amateur and professional. Even if it 


VOLUME 40, NUMBER 3 137 


should not prove optimal for mass rearing of a given species, our diet 
may facilitate culturing of species not otherwise culturable, especially 


in the field. 


ACKNOWLEDGMENTS 


This study was supported by the Deutsche Forschungsgemeinschaft (SFB4/B6). We 
thank M. Bairov for laboratory assistance, A. Egelhaaf for partly providing rearing fa- 
cilities, and A. Watson for identification of several species. We are also indebted to D. 
Schneider, to P. M. Brakefield, and, especially, to A. W. R. McCrae for valuable com- 
ments on the manuscript. Suggestions by an anonymous reviewer helped to streamline 
the manuscript, and are acknowledged with thanks. 


LITERATURE CITED 


BATHON, H. 1977. Die Zucht des Barenspinners Spilosoma maculosa Stoll. (Lep. Arc- 
tiidae) auf einem kiinstlichen Nahrmedium. Ber. Offb. Ver. Naturkde. 80:54-60. 

BELL, R. A. & F. G. JOACHIM. 1976. Techniques for rearing laboratory colonies of 
tobacco hornworms and pink bollworms. Ann. Entomol. Soc. Am. 69:365-373. 

CONNER, W. E., T. EISNER, R. K. VANDER MEER, A. GUERRERO & J. MEINWALD. 1981. 
Precopulatory sexual interaction in an arctiid moth (Utetheisa ornatrix): Role of a 
pheromone derived from dietary alkaloids. Behav. Ecol. Sociobiol. 9:227-235. 

EL-GuINpDy, M. A., M. M. EL-SAYED & Y. H. Issa. 1979. Biological and toxicological 
studies on the cotton leafworm Spodoptera littoralis Boisd. reared on natural and 
artificial diets. Z. PflKrankh. 86(3/4):180-189. 

GOTTEL, M. S. & B. J. R. PHILOGENE. 1978. Laboratory rearing of the banded wooly- 
bear, Pyrrharctia (Isia) isabella (Lep.: Arctiidae), on different diets with notes on 
the biology of the species. Can. Entomol. 110:1077-1086. 

Hiral, K. 1976. A simple artificial diet for mass rearing of the armyworm, Leucania 
separata Walker (Lep.: Noctuidae). Appl. Entomol. Zool. 11:278-283. 

McGInnis, A. J. & R. KAsTING. 1964. Comparison of gravimetric and chromic oxide 
methods for measuring percentage utilization and consumption of food by phytoph- 
agous insects. J. Insect Physiol. 10:984—995. 

Moreau, J. P. 1965. A propos de la biologie d’Arctia caja L. (Lep.: Arctiidae). Proc. 
12th Int. Congr. Entomol. Lond. (1964):539. 

SCHLIEPER, C. A. 1982. Grundfragen der Ernahrung. Vlg. Handwerk und Technik, 
Hamburg. 421 pp. 

SHOREY, H. H. & R. L. HALE. 1965. Mass-rearing of the larvae of nine noctuid species 
on a simple artificial medium. J. Econ. Entomol. 58:522-524. 

SINGH, P. 1977. Artificial diets for insects, mites and spiders. IFI/Plenum Data Com- 
pany, New York. 594 pp. 

SINGH, P. & R. F. Moore. 1985. Handbook of insect rearing (Vol. II). Elsevier Science 
Publishers, Amsterdam. 

VAIL, P. V., T. J. HENNEBERRY & R. PENGALDEN. 1967. An artificial diet for rearing 
the salt marsh caterpillar, Estigmene acrea (Lep.: Arctiidae), with notes on the 
biology of the species. Ann. Entomol. Soc. Am. 60:134-188. 

VANDERZANT, E. S. 1967. Wheat-germ diets for insects: rearing the boll weevil and 
the saltmarsh caterpillar. Ann. Entomol. Soc. Am. 60:1062-1066. 

1974. Development, significance, and application of artificial diets for insects. 

Ann. Rev. Entomol. 19:189-160. 





Received for publication 14 January 1986; accepted 6 June 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 138-157 


EVOLUTION AND IDENTIFICATION OF THE NEW WORLD 
HAIRSTREAK BUTTERFLIES (LYCAENIDAE: EUMAEINI): 
ELIOT’S TRICHONIS SECTION AND 
TRICHONIS HEWITSON 


ROBERT K. ROBBINS 


Department of Entomology, MRC NHB 127, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560 


ABSTRACT. I revise the lycaenid genus Trichonis Hewitson (Theclinae: Eumaeini), 
establish the new combination Trichonis hyacinthus Cramer, and make Papilio thenaus 
Cramer 1777 a junior synonym of P. hyacinthus Cramer 1775. The genus consists of 
two species, T. hyacinthus and T. immaculata, which differ in wing pattern, forewing 
shape, color of androconial scales, and length of the third palpal segment. I then assess 
Eliot’s higher classification of the Eumaeini. I examine leg, genitalia, and wing mor- 
phology, and conclude that Eliot’s Trichonis Section is diphyletic. The unusual male 
foretarsus of Trichonis appears to have evolved independently three times in the Eu- 
maeini. 


Eliot (1978) and Clench (1964, pers. comm. 1978) proposed different 
higher classifications for the “New World hairstreaks’ (Lycaenidae: 
Theclinae: Eumaeini). Eliot divided them into a Trichonis Section— 
for genera Trichonis Hewitson and Micandra Schatz—and an enor- 
mous Eumaeus Section—for the remaining genera (64 available 
names). Clench also divided these butterflies in two groups; in one he 
isolated Eumaeus Hubner (his Eumaeini), and in the other he lumped 
the remaining genera (his Strymonini). 

The purpose of this paper is to assess the evidence for Eliot’s pro- 
visional higher classification of the Eumaeini. Specifically, I consider 
whether the Trichonis Section is a monophyletic group. As basic in- 
formation needed to answer this question, I revise Trichonis. Mican- 
dra, which Clench (1971) treated preliminarily, needed little work for 
the purpose of this paper. I then discuss the evidence for Eliot’s clas- 
sification. 


GENUS TRICHONIS 


Trichonis consists of two species known only from males, T. theanus 
Cramer (Fig. la, b) and T. immaculata Lathy (Fig. 2a, b). In 1865, 
Hewitson named Trichonis, primarily on the basis of an unusual male 
foretarsus in T. theanus, which he described as “‘exarticulate, robust, 
and broad beyond the middle” (Fig. 7a, b). Hewitson also illustrated 
the “female” of T. thenaus, and described its foretarsus as “of the 
usual form, jointed and spined.”’ Lathy (1930) pointed out that Hew- 
itson’s female was the male of a second species, which he named T. 
immaculata. Lathy did not, however, compare foreleg morphology of 


VOLUME 40, NUMBER 3 139 





Fic. 1. Adult Trichonis hyacinthus. (a) male upperside, (b) male underside, (c) fe- 
male upperside, (d) female underside. 


T. theanus and T. immaculata, nor did he discuss whether Hewitson’s 
characterization of Trichonis was valid. No females have been asso- 
ciated with either species. 

I propose that Papilio hyacinthus Cramer (Fig. lc, d)—Cramer 
named all butterflies in Papilio—and a second phenotypically similar 
species (Fig. 2c, d) are the females of T. theanus and T. immaculata. 
Neither “female species’ has been associated with males, but both 
share with T. theanus and T. immaculata a pastel blue or blue-green 
color on the frons and ventral wings that is unique among the Eumae- 
ini. Both sexes have truncate forewings, and share similar geographical 
distributions. Further, genital morphology, which is discussed more 
fully later, indicates that both males and females belong to a similar 
section of the Eumaeini. 

I associate female T. hyacinthus with male T. theanus and the new 
female with male T. immaculata. The bases for this action are distri- 
bution (Fig. 3) and forewing shape (Fig. 4). Briefly, T. hyacinthus and 
T. theanus are known only from the Guianas and Lower Amazon while 
the new female and T. immaculata occur there and in the Upper 
Amazon; the new female and a male of T. immaculata were collected 


140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Adult Trichonis immaculata. (a) male upperside, (b) male underside, (c) 
female upperside, (d) female underside. 


at the same locality in northern Peru. As detailed below, the forewing 
apex is more truncate in male T. immaculata than in male T. theanus 
and in the new female than in female T. hyacinthus. 

Characters unique to Trichonis include a pastel blue or blue-green 
frons and ventral ground color, and the genitalia, whose structure (Figs. 
5 & 6) differs quantitatively from other eumaeines. I do not know 
which species or species group is its closest relative. Trichonis may be 
enlarged as our understanding of eumaeine phylogeny increases. 


Key to Trichonis Species and Sexes 


1. Ventral hindwings without transverse brown lines (Fig. 1b, 2b) ................ (males)... 2 
Ventral hindwings with transverse brown lines (Fig. 1d, 2d) (females) ... 3 

2. Inner edge of dorsal forewing marginal band “smooth” (Fig. 1a), not scalloped. 

Ventral forewing androconial patch almost touching upper part of discal cell 
(Big) Aa jie ou sah Ne lot a nS Ae ele a ce male hyacinthus 

Inner edge of dorsal forewing marginal band scalloped (Fig. 2a). Ventral forewing 
androconial patch barely enters discal cell (Fig. 4c) 000 eccco male immaculata 

3. Ventral hindwing with a white band between major brown transverse lines (Fig. 
Id); hindwingewith.a\ tail)... eee ees ee female hyacinthus 

Ventral hindwing with a blue or blue-green (ground color) band between major 
brown transverse lines (Fig. 2d); hindwing without a tail _... female immaculata 


VOLUME 40, NUMBER 3 141 


TRICHONIS SYSTEMATICS 


Nomenclature 


Trichonis Hewitson (1865): Hewitson described Trichonis in the 
Lycaenidae with Cramer’s Papilio theanus as the only species in the 
genus. It is the type species by monotypy (Hemming 1967). Lathy 
(1930) subsequently added T. immaculata. 

Papilio hyacinthus Cramer (1775): Cramer described Papilio hy- 
acinthus from the West Indies. He included a brief description and a 
figure of the underside, but did not mention its sex. Various authors 
(Fabricius 1782, Butler 1870, Draudt 1919-1920) discussed it, but their 
text paraphrased the original description, and illustrations were copies 
of Cramer’s (sometimes poorly done, such as Seitz [Draudt 1919-1920)}). 
No one has mentioned P. hyacinthus for more than half a century 
except to note that it is unknown from the West Indies (Comstock & 
Huntington 1943). 

There is no other species with which the figure of P. hyacinthus can 
be confused; the white band sandwiched between two transverse brown 
lines on the ventral hindwing is distinctive. All known specimens are 
females. 

There are no potential lectotypes in the Artis Collection, Instituut 
voor Taxonomische Zodlogie, Zodlogisch Museum, Universiteit van 
Amsterdam (Hogenes, pers. comm.), at the Rijksmuseum van Natuur- 
liike Historie in Leiden (de Jong, pers. comm.), or in the British Mu- 
seum (Natural History) (BMNH). However, identification of Cramer’s 
P. hyacinthus poses no problems, and a type is not needed. Trichonis 
hyacinthus is a New Combination. 

Papilio theanus Cramer (1777): Cramer described Papilio theanus 
from Surinam with a brief description and a figure of the ventral 
surface, on which “‘androconial”’ patches are evident. Hewitson (1862- 
1878) illustrated the male and female, but Lathy (1930) noted that 
Hewitson’s female is the male of a second species (which he named 
Trichonis immaculata). All known specimens are males. 

Identification of P. theanus poses a problem. Males of both Trich- 
onis species have extremely similar ventral wing patterns (Fig. 1b, 2b), 
and Cramer illustrated only the ventral surface. The species can be 
distinguished, however, by forewing shape and extent of the “polished 
spot” surrounding the forewing androconia. Unfortunately, the fore- 
wing shape in Cramer’s illustration is inaccurate (the curvature at the 
forewing apex is too great for either species). The polished spot in the 
original illustration (in the BMNH library) extends almost to the radial 
vein, which is not the case in the few specimens of T. immaculata that 
I examined. However, considering the inaccuracy of other characters 


142 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 















































b KS 
West Longitude 











Oblique Conic Conformal Projection Smithsonian Institution 1983 
Prepoted by Theophilus Brit! Griswold 


Fic. 3. Distribution of Trichonis. Solid dots designate exact localities, hollow dots 
represent generalized localities such as “Surinam” or “Maranham.” (a) T. hyacinthus, 
(b) T. immaculata. 


‘ 
= 3 
= ~ 
~ 

iS 

5 

3 

=> 

= 

Es 

a 

Oo 

E 

oO 

a 

2 

“a 

3 

= 

~ 

= 

— 

13) 

Q TG "S6 

2S 

8 

NS 

os 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in the original figure, I hesitate to base a specific identification on so 
minute a detail. A type specimen is desirable. 

I designate as lectotype of P. thenaus a specimen in the Artis Col- 
lection (Zodlogisch Museum, Amsterdam). This specimen fits Cramer’s 
original figure well, and was probably seen by Cramer even if it was 
not the model for his figure. There are no potential lectotypes in Leiden 
or London. The specimen bears two labels: one with the number “16” 
and one with the text ““Trichonis Theanus Cram,” written in the hand 
of Snellen, according to Willem Hogenes, Keeper of Lepidoptera at the 
Zoologisch Museum. I have added a black-bordered label which reads: 
“Lectotype Papilio theanus Cramer, 1777; by R. Robbins.”” The words 
“Lectotype” and “by” are printed in red and the remainder of the 
label is hand-written in black ink. This lectotype designation will pre- 
vent confusion in the future, and maintain the previous identifications 
of P. theanus used by Hewitson (1862-1878), Staudinger (1884-1888), 
Schatz and Rober (1885-1892), and Lathy (1930). 

I already outlined the evidence for considering P. theanus to be the 
male of P. hyacinthus, and now designate P. theanus Cramer a junior 
synonym of P. hyacinthus Cramer; New Synonymy. 

Trichonis immaculata Lathy (1930): Lathy described T. immacu- 
lata from two males: one without locality data (presumably in the 
Museum National D’Histoire Naturelle) and one which was the model 
for Hewitson’s (1862-1878) “female” illustration of T. theanus 
(BMNH). Either can eventually be designated a lectotype, but identi- 
fication poses no problems. Male T. immaculata can be distinguished 
from male T. hyacinthus (=T. theanus) by the “smooth” dorsal fore- 
wing border, as noted in the above key. 


Geographic Distribution 


Trichonis occurs in the Guianas and throughout the lowland Amazon 
Valley from the mouth of the Amazon River at the Atlantic Ocean to 
the headwaters at the base of the eastern Andes (Fig. 3). The genus is 
unrecorded from the West Indies except for Cramer’s unverified T. 
hyacinthus record. 

I examined the following specimens of T. hyacinthus (Fig. 3a) in 
the BMNH except where noted. Guyana (formerly British Guiana)— 
5 males; Surinam—2 males; French Guiana (sometimes labelled Cay- 
enne, which is thus inseparable from the present day city of that 
name)—5 males and 2 females, Maroni—1 female, Maroni R., St. Lau- 
rent—1 male; Brasil, Para—8 males and 1 female (1 male and 1 female 
in NMNH—National Museum of Natural History, Smithsonian Insti- 
tution), Maranham—1 female; No Data—2 males from the Felder 


VOLUME 40, NUMBER 3 145 





Fic. 5. Trichonis male genitalia. From left to right: dorsal, lateral, and ventral views. 
(a) T. hyacinthus, (b) T. immaculata. Scale line is 1 mm. 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 6. Trichonis female genitalia, ventral view of bursa copulatrix. (a) T. hyacin- 
thus, (b) T. immaculata. Scale line is 1 mm. 


Collection, 1 male (lectotype) in Zodlogisch Museum. None of the spec- 
imens has a date of capture. 

I saw the following specimens of T. immaculata (Fig. 3b) in the 
BMNH unless otherwise noted. Surinam—1 male, Paramaribo—1 male 
(NMNH); French Guiana—(labelled Cayenne, Hewitson’s female fig- 
ure)—1 male, Gourdonville, R. Kourov—1 male; Brasil, Amazonas, 
Manacapuru—1 male (Carnegie Museum of Natural History); Peru— 
Loreto, Iquitos, Rio Cachiyaca (usually spelled Cachiyacu)—1 male 
and 1 female, Madre de Dios, 30 km SW of Pto. Maldonado—1 male 
(private collection of Dan Bogar). The Paramaribo specimen was col- 
lected in September /October, the Manacapuru specimen in May 1926, 
and the Madre de Dios specimen on 22 October 1983 at 0930 h. 





Morphology 


Antennae: 40-44 segments. The beginning of the club is not clearly 
defined; rather, the segments gradually increase in size and become 
slightly flattened (dried specimens) on either side. As a result, the club 


VOLUME 40, NUMBER 3 147 


segments are elliptically shaped. The club is composed of 18 or 19 
segments. The nudum (area without scales) is found on the 5 apical 
segments dorsally and on the 23-24 apical segments ventrally. There 
are incomplete white annulations around the segments on the stalk and 
the first few segments of the club, where the white scaling sometimes 
coalesces into a short line. The few specimens with intact antennae 
reveal no difference between the sexes or between the species. 

Eyes: The eyes have short sparse hairs, and are slightly emarginate 
at the antennal bases. Hewitson (1862-1868) and Draudt (1919-1920) 
incorrectly reported the eyes as smooth, which, if true, would have 
made Trichonis unique among the Eumaeini (Eliot 19783). There is a 
ring of scales (Some white and some blue) surrounding the eyes, but 
interrupted by the antennal scape and chaetosema. 

Frons: The frons is covered with downward oriented blue scales 
lined laterally with white scales. 

Labial Palps: I measured length of the third palpal segment (with 
an ocular scale) because it appeared to be sexually dimorphic and to 
differ interspecifically. The male T. hyacinthus third palpal segment 
(mean = 0.31 mm, SD = 0.026, N = 4) is significantly shorter than the 
female segment (mean = 0.83, SD = 0.193, N = 8; P < 0.05, t-test for 
unequal variances, Sokal & Rohlf 1969). Likewise, this length is sig- 
nificantly shorter in male T. immaculata (male: mean = 0.40, SD = 
0.011, N =38; female: mean = 0.78, N = 1; P < 0.01, t-test for one ob- 
servation with mean of a sample). Although such sexual dimorphism 
has apparently not been reported for eumaeines, a quick survey of 
other genera indicated that it occurs frequently. Length of the third 
segment is also significantly longer in male T. immaculata than in 
male T. hyacinthus (P < 0.01, t-test), contrary to Hewitson’s (1862— 
1878) claim that they are the same. There is no evidence that the 
females are different (P > 0.05, t-test). 

Thorax: Thorax, legs, and wings are covered with blue or blue-green 
scales. The color varies individually, not seeming to be species specific, 
except that the dorsal wing color of male T. hyacinthus is consistently 
more greenish than that of male T. immaculata. 

Legs: Discussed later in the section on phylogenetic affinities. 

Wing Venation: Schatz and Rober (1885-1892) figured the venation 
of male T. hyacinthus (as male T. theanus). The venation of both 
species and sexes (Fig. 4) is typical of the Eumaeini with 10 forewing 
veins. The position of forewing veins R, and M, varies interspecifically 
among eumaeine species. In Trichonis, forewing vein R, arises from 
the discal cell, and forewing vein M, arises slightly nearer vein M, 
than M,. 

Wing Shape: The forewing apex is strongly truncated in both 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


species—more so in males than in females and more so in T. immac- 
ulata than in T. hyacinthus (Fig. 4). To show this, I measured the 
angle between a line connecting the ends of veins R, and R, on the 
costa and a line connecting the ends of veins M, and M, on the outer 
margin. The angle of male T. hyacinthus was at or slightly more acute 
than 90° (N = 4) while male T. immaculata was always more than 100° 
(N = 4). Female T. hyacinthus ranged from 75° to 83° (N = 3). I mea- 
sured the forewing angle of the single T. immaculata female at 87°. 

Female T. hyacinthus is tailed at hindwing vein Cu, (Fig. 4b), but 
neither its male nor either sex of T. immaculata has an indication of 
a tail (Fig. 4a, ec, d). 

Size: | measured forewing length from the base of the radial vein to 
the forewing apex as follows: male T. hyacinthus, mean = 1.7 cm, 
SD = 0.17, N =4; female T. hyacinthus, mean = 1.8 cm, SD = 0.416, 
N = 3; male T. immaculata, mean = 1.5 cm, SD = 0.05, N = 4; female 
T. immaculata, mean = 1.5 cm, N = 1. 

Androconia: The venation drawings show the position and outline 
of the androconial patches on the ventral forewing and dorsal hindwing 
(Fig. 4). Each patch is composed of two or three parts. Forewing and 
hindwing inner patches of T. hyacinthus are dark brown while those 
of T. immaculata are beige. Around the inner patches, and contrasting 
with them, is an area of silver scales which give the impression of being 
a “polished spot.” The extent of the polished area is poorly defined on 
some parts of the wings, as shown by the trailing dotted lines in the 
figures. The polished area immediately surrounding the inner patches 
has a greenish tint, but once again, this area is poorly defined. The 
extent of the polished area on the ventral forewing differs in the two 
species. In T. hyacinthus it extends through the discal cell, and touches 
or nearly touches the radial vein at the top of the cell. In T. immac- 
ulata the polished spot extends less than half way through the cell. 
(Figs. 1b, 2b do not show this difference clearly unless used in con- 
junction with Fig. 4.) Eliot (1973) published an outline drawing of a 
hindwing androconium in T. hyacinthus. 

Male Genitalia (Fig. 5): Saccus almost lacking, vinculum thick ven- 
trally, valvae small, penis thick with a single terminal cornutus. I found 
no consistent differences between the species. The specimens illustrated 
in Fig. 5 represent the extremes of individual genital variation in shape 
of the valvae and ventral vinculum, and in the position of the vinculum 
strut. 

Female Genitalia (Fig. 6): Ductus bursae short and sclerotized, con- 
cave dorsally for its entire length and twisted dextrally (not evident 
from the figure). The corpus bursae is exceedingly long compared to 
the ductus bursae, and is posteriorly constricted and lightly sclerotized. 


VOLUME 40, NUMBER 3 149 


The ductus seminalis, which arises from the posterior end of the corpus 
bursae, is “off-center” to the right side of the female. There are a pair 
of signa as illustrated. As with the males, there are no evident differ- 
ences between the species. 


Biology 


Almost nothing is known about the biology of Trichonis. Since most 
of the known specimens were collected long ago, I speculate that Trich- 
onis species inhabit primary forest. Because of extensive deforestation, 
modern visitors to the Amazon Basin rarely collect in virgin jungle. 
Indeed, the one recent collection of T. immaculata was in the Tam- 
bopata Reserve (Madre de Dios, Peru), where the jungle is protected 
from cutting. 


Similar Species 


The wing pattern of male Trichonis is so distinctive that it cannot 
be confused with that of species in other genera. Female Trichonis, 
however, are superficially similar to, and might be confused with, 
“Thecla” tagyra Hewitson and “Thecla”’ floralia Druce (which I con- 
sider a senior synonym of “Thecla” tagyroides Lathy). “Thecla”’ ta- 
gyra and “T.”’ floralia have a light blue frons and ventral ground color 
that is similar to Trichonis, but of a different quality when compared 
side by side. They are most easily differentiated from Trichonis by 
two superficial characters: they possess a red anal lobe on the dorsal 
hindwing, and black transverse lines on the ventral hindwing. Trich- 
onis females lack a red anal lobe and have brown transverse lines on 
the ventral hindwing. I tentatively place tagyra and floralia in Evenus 
Hubner, a genus I consider unrelated to Trichonis on the basis of 
androconial structure and genital morphology. 


PHYLOGENETIC AFFINITIES 


Eliot (1978) placed Trichonis in the Eumaeini because it shares the 
diagnostic characters of the tribe: 10 forewing veins, “greyhound- 
shaped” male genitalia lacking a juxta, a stubby-tipped male foretarsus 
(at least in T. immaculata), and hairy eyes. I address the question of 
its phylogenetic affinities within the Eumaeini by discussing leg mor- 
phology, genitalia, and wing structures. 


Legs 
The lycaenid male foretarsus is unique among the Lepidoptera. The 
tarsomeres are fused into one segment, lack tarsal claws, are used for 
walking, and possess on the ventral surface “smooth-walled sensilla” 
with an opening at the tip and ‘“‘spines’—presumed sensilla with 


150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


longitudinal striations and no opening at the tip (Fig. 8). Although 
male Riodinidae, Libytheidae, and Nymphalidae also have clawless 
foretarsi, they lack spines and smooth-walled sensilla on the foretarsus, 
and do not use their forelegs for walking. Some male lycaenids have a 
segmented and clawed foretarsus (Eliot [1973] lists genera), but evi- 
dently this structure has been independently re-expressed a number of 
times within the Lycaenidae (Eliot 1978, Robbins, in prep.). The male 
lycaenid foretarsus may be stubby-tipped (characteristic of the Eu- 
maeini) or tapered to a sharp point (Clench 1955, Eliot 1973). 

The male foretarus of T. hyacinthus is different from that of T. 
immaculata. The male of T. immaculata has a typical eumaeine fore- 
tarsus (Fig. 7c, d); it is cylindrical and stubby-tipped, and possesses 
spines and smooth-walled sensilla. Although Hewitson (1862-1878) 
claimed that it was a normal female foreleg, it lacks the claws and 
segmentation that occur in all lycaenid females. Unlike that of T. 
immaculata, the foretarsus of male T. hyacinthus (Figs. 7a, b, 8a, b), 
has a mid-ventral bulge (Hewitson 1862-1878), spines only at the tip 
(Eliot 1973), and smaller, somewhat flattened spinelike projections cov- 
ering the ventral surface except for the tip. The spinelike projections, 
however, are striated like normal eumaeine spines (Fig. 8b), and there 
is a sharp transitional area of intermediate-sized spines (Fig. 8a). On 
the basis of this observation, I consider the spinelike projections to be 
small spines. Despite its unusual morphology, the male foretarsus of T. 
hyacinthus is technically lycaenid in that it is fused and possesses spines 
and smooth-walled sensilla (Fig. 8a). 

Two other eumaeines besides T. hyacinthus have a stout, centrally 
swollen foretarsus spined only at the tip. The first is Micandra platyp- 
tera Felder & Felder (Figs. 7e, f, 8c, d), as Eliot (1973) noted. The 
foretarsus, however, lacks most of the mid-ventral bulge, and the tran- 
sition in spine size is more gradual than in T. hyacinthus. The second 
species is “Thecla”’ myrtusa Hewitson (Figs. 7i, j, 8e, f). Its foretarsus 
is shaped differently than the other two, has fewer long spines at the 
tip, and, most notably, the small spines occur primarily on the inner 
face of foretarsus (not evident from the figures). 

Micandra, like Trichonis, contains only one species with an atypical 
male foretarsus. Clench (1971) placed playtptera and tongida Clench 
in Micandra on the basis of venation, genitalia, and wing pattern, and 
listed eight other potential member species. I examined the male gen- 
italia and venation of ion Druce, comae Druce, cyda Godman & Sal- 
vin, aegides Felder & Felder, and amplitudo Druce (probably a syn- 
onym of aegides), and all belong to Micandra as Clench characterized 
it. The “‘invaginated pocket’ that Clench described on the male genital 
valvae is actually a process pointing caudally. Also, all species have 


VOLUME 40, NUMBER 3 


Fic. 7. Male forelegs. Lateral view on left and ventral view on right. (a & b) T. 
hyacinthus, 35x, (c & d) T. immaculata, 40x, (e & f) Micandra platyptera, 24x, (g & 
h) M. comae, 37x, (i & j) “Thecla” myrtusa — lateral view is of outer surface, two 


> 


spines at tip of foretarsus are broken, 32x, (k & 1) “Thecla” myrtea, 44x. 


dorsal forewing androconia, contrary to Clench’s key. Except for M. 
platyptera, however, these species have regularly spined male foretarsi 
lacking a ventral bulge (Fig. 7g, h). 





JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


2 gm ST Si Dee 


~I YX SII J eS 


Fic. 8. Male forelegs. Ventral view. (a) T. hyacinthus, distal end, 110x, (b) T. 
hyacinthus, small flattened spines with striations, 525 =, (c) Micandra platyptera, distal 
end, 85x, r—regular lycaenid “spine,” p—small, flattened “spine,” s—“smooth-walled 
sensillum,” (d) M. platyptera, transition between small and regular spines, 210%x, (e) 
“Thecla” myrtusa, distal end with inner face on top, 190x, (f) “T.” myrtusa, small 
spines, 530. 


The species most closely related to “Thecla” myrtusa also lack a 
centrally swollen foretarsus spined only at the tip. Although phyloge- 
netic relations are not yet worked out, “T.” myrtusa appears to be 
most closely related to two groups of hairstreaks. The first group con- 





VOLUME 40, NUMBER 3 158 


tains “Thecla”’ myrtea Hewitson, “T.” falerina Hewitson, “T.” eunus 
Godman & Salvin, and “T.” thara Hewitson, and is defined by a unique 
dorsal hindwing “‘androconial’’ patch in which the scales have co- 
alesced to form a thin foil-like lamination on the wing membrane. 
“Thecla” myrtusa shares with them a dark androconial patch at the 
base of the ventral forewing cubital vein, and shares a similar ventral 
wing pattern with “T.” myrtea and “T.” falerina. The second poten- 
tial “closest relative” of “T.’’ myrtusa is Allosmaitia Clench. It shares 
with “T.’’ myrtusa beige (gray in some specimens) dorsal forewing 
androconia interspersed with regular wing scales, a character that 
Clench (1964) overlooked in Allosmaitia. The species in the “T.” myr- 
tea group and in Allosmaitia have male foretarsi with regular rows of 
spines and without a central bulge (Fig. 7k, l). 

“Short spines’ are currently known only on the foretarsi of T. hy- 
acinthus, M. platyptera, and “T.”’ myrtusa, but are difficult to see 
under a binocular microscope. Since I did not look at the male foretarsi 
of all their relatives under greater magnification, it is possible that some 
have short spines interspersed with regular ones. 


Genitalia 


The genitalia of Trichonis are quantitatively distinct from those of 
other eumaeines, and lack unusual qualitative characters that might 
be shared with other genera. Thus, in this case genital structures give, 
at best, an imprecise indication of relationship. 

I found two major patterns of correlated genital structures among 
eumaeines. The first is characterized by a thick ventral vinculum, stout 
penis, taut manica (the membrane attaching the penis to the valvae) 
allowing little penial movement, no ventral processes on the lateral 
tegumen, and short ductus bursae with a simple cervix (the anterior 
ductus bursae ends abruptly with almost no change in structure). The 
second pattern is the antithesis of the first: a thin ventral vinculum, 
thin penis, loose manica, ventral processes of the tegumen present, and 
long ductus bursae (usually as long as the corpus bursae) with a ““com- 
plex” cervix in which the shape of the anterior ductus bursae is dif- 
ferent from the remainder of the ductus bursae. Examples of the first 
pattern are Parrhasius Hubner, Iaspis Kaye, Erora Scudder, and Sym- 
biopsis Nicolay, and of the second, Mithras Hiibner, Evenus, Theritas 
Hubner, and Rekoa Kaye. The two patterns represent extreme modes 
along a continuum of genital patterns, so that many species are inter- 
mediate and congenors may differ in one or two of these characters. 
However, Trichonis fits the first pattern while Micandra and “‘T.” 
myrtusa fit the second. In the absence of qualitative characters, this 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


evidence indicates that Trichonis is not phylogenetically close to Mi- 
candra and “T.” myrtusa. 


Wing Venation, Shape, Pattern, and Androconia 


Trichonis wing venation is commonplace, but the truncate forewing 
is unusual. Although various other eumaeine species, such as Pan- 
thiades bitias Cramer, have truncate forewings, there are no other 
shared characters to support a close relation with Trichonis. Species in 
the “Thecla” rocena Hewitson complex, however, share a relatively 
rectangular forewing shape and ventral forewing androconia with 
Trichonis, and have broadly similar genitalia (though with many points 
of difference). The coxa, femur, and tibia of “T.”’ rocena male forelegs 
are abnormal, but the tarsus, unlike T. hyacinthus, is normal. “Thecla”’ 
rocena and allies may be close relatives of Trichonis, but I did not 
find definitive evidence supporting this relation. 

Wing venation may be a good indicator of Micandra’s relations. 
Both Schatz and Roéber (1885-1892) and Eliot (1973) illustrated the 
unusual distal forewing discal cell venation, which is shared with minor 
variation by all species of Micandra. Species in other genera, such as 
“Thecla” timaeus Felder & Felder and relatives, also share this char- 
acter, and are undoubtedly close relatives of Micandra. 

Schatz and Rober (1885-1892) and Clench (1971) noted that Mi- 
candra forewing vein R, originates far basad of the other radials, and 
is situated next to Sc for most of its length. This distinctive venation is 
found in Micandra, “Thecla” timaeus, “Thecla” eronos Druce and 
relatives, ““Thecla’”’ auda Hewitson and relatives, and “Thecla” busa 
Godman & Salvin and relatives, but does not occur in either Trichonis 
or “Thecla’”’ myrtusa. I consider it likely that this character state will 
eventually characterize a monophyletic assemblage of eumaeine gen- 
era, and if so, indicates that the closest relatives of Micandra are not 
Trichonis or “T.”’ myrtusa. 

The ventral wing pattern of Trichonis is unique, and does not pro- 
vide clues to its systematic position. I mentioned earlier that I consider 
the superficial similarity between the ventral wing patterns of female 
Trichonis and two species of Evenus to be convergence. 

Male Trichonis androconia on the dorsal hindwing and ventral fore- 
wing, located where the wings overlap, are also of no help in working 
out phylogenetic position. Many species scattered throughout the Eu- 
maeini, as well as the Deuodorigini, a close relative of the Eumaeini 
(Eliot 1973), have androconia where the wings overlap. The exact 
structure of Trichonis androconial patches, as detailed above, is unique, 
so far as I am aware. 

Eliot (1973) supported his Trichonis Section with the observation 


VOLUME 40, NUMBER 3 155 


that Trichonis and Micandra androconia are the same size or larger 
than ordinary scales—in contrast to the “small” androconia of the Eu- 
maeus Section. My results do not support this observation. I found that 
the dorsal forewing androconia of “Thecla’”’ mycon Godman & Salvin 
average 2.8 times larger than adjacent iridescent blue scales (N = 10). 
Likewise, the dorsal forewing distal androconia of Atlides halesus Cra- 
mer average 1.5 times larger than dorsal forewing blue scales (N = 
10). Further, Eliot (1973:402) listed other species with ventral forewing 
androconia that are larger than “‘ordinary” scales. A quick survey in- 
dicated that this character state is widespread in the tribe. Further, 
“ordinary” wing scales can vary in size by a factor of 7 (Gray 1962). 
I doubt that relative androconia size will be a useful character state. 


Conclusions 


There are three evolutionary hypotheses that might account for the 
information just presented. The first hypothesis is that T. hyacinthus, 
M. platyptera, and “T.”’ myrtusa form a monophyletic group defined 
by their male foretarsus. Consistent with this hypothesis is the obser- 
vation that males of the first two species have round hindwings lacking 
tails while their females are tailed. However, wing pattern, genital, 
androconial, and venational characters are inconsistent with this hy- 
pothesis, and indicate that T. hyacinthus is congeneric with T. im- 
maculata, M. platyptera with the species that Clench (1971) placed in 
Micandra, and “‘T.” myrtusa with the “T.” myrtea group and/or 
Allosmaitia. Further, sexual dimorphism in the tailed condition occurs 
in other eumaeines with typical eumaeine male foretarsi, such as Erora 
phrosine Druce and “Thecla”’ timaeus. 

The second hypothesis is that Trichonis, Micandra, and the eventual 
generic assignment of “Thecla’”’ myrtusa form a monophyletic group 
defined by the tendency to express the atypical male foretarsus. This 
group would approximately correspond to Eliot’s (1978) Trichonis Sec- 
tion. However, there are no similarities in genitalia, venation, wing 
pattern, or androconia to support this hypothesis. Even the unusual 
tailed dimorphism mentioned in the previous paragraph occurs in only 
one species of Trichonis and one species of Micandra. Further, genital 
and venational structures, as discussed in the previous section, indicate 
that Trichonis belongs to a different group of eumaeine genera than 
Micandra and “‘T.”’ myrtusa. 

The third hypothesis is that the atypical male foretarsus of T. hy- 
acinthus, M. platyptera, and “T.” myrtusa has evolved independently 
three times. The distribution of other character states is consistent with 
this hypothesis, and indicates that Eliot’s Trichonis Section is diphy- 
letic. 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


An objection to this conclusion is that the repeated evolution of a 
qualitatively distinct foreleg is unlikely. However, the three atypical 
forelegs are not identical, casting doubt on their homology. Further, if 
a slight change during development of the male foreleg were respon- 
sible for the atypical foretarsus, then only a small genetic change, such 
as a mutation in a promoter or in the timing of transcription, is nec- 
essary to explain its repeated independent occurrence. Indeed, eukary- 
otic genes that regulate timing of development are now known (Am- 
bros & Horvitz 1984). As the regulatory mechanisms determining 
development of insect leg structures are worked out, as they are being 
done for egg chorion structures (reviewed in Kafatos 1981) and wing 


pattern (Nijhout 1978, 1980a, b, 1981, 1984), it will be possible to test 
this idea. 


ACKNOWLEDGMENTS 


I thank R. I. Vane-Wright, P. Ackery, R. Smiles (British Museum [Natural History]), 
C. W. Young (Carnegie Museum of Natural History), and D. Bogar (Harrisburg, PA) for 
allowing me access to Trichonis specimens in collections under their care. I thank R. de 
Jong for good natured correspondence on Cramer types in Leiden, and particularly thank 
Willem Hogenes, Keeper of Lepidoptera at the Zodlogisch Museum, for loaning me the 
lectotype of T. theanus, and for sending me historical information on Cramer types in 
the Artis Collection. I thank S. S. Nicolay for providing me with unpublished information 
on eumaeine morphology. I thank J. M. Burns, J. N. Eliot, J. S. Glassberg, S. S. Nicolay, 
and R. W. Poole for reading and commenting on the manuscript. 


LITERATURE CITED 


AMBROS, V. & H. R. Horvitz. 1984. Heterochronic mutants of the nematode Caeno- 
rhabditis elegans. Science 226:409-416. 

BUTLER, A. G. 1870. Catalogue of diurnal Lepidoptera described by Fabricius in the 
collection of the British Museum. London. 303 pp. + 8 pl. 

CLENCH, H. K. 1955. Revised classification of the butterfly family Lycaenidae and its 
allies. Ann. Carnegie Mus. 33:261-274. 

1964. A synopsis of the West Indian Lycaenidae, with remarks on their zoo- 

geography. J. Res. Lepid. 2:247-270. 

1971. Two new hairstreaks from Mexico (Lepidoptera: Lycaenidae). Bull. Allyn 
Mus. No. 3. 6 pp. 

Comstock, W. P. & E. I. HUNTINGTON. 1943. Lycaenidae of the Antilles (Lepidoptera, 
Rhopalocera). Ann. N.Y. Acad. Sci. 45:49-130. 

CRAMER, P. 1775-1776. De Uitlandsche Kapellen voorkomende in de drie Waereld- 
deelen Asia, Africa en America. S. J. Baalde, Amsterdam. Vol. I. 156 pp., 96 pl. 
1777. De Uitlandsche Kapellen voorkomende in de drie Waereld-deelen Asia, 

Africa en America. S. J. Baalde, Amsterdam. Vol. II. 151 pp., 96 pl. 

DraAupDT, M. 1919-1920. Thecla F. In Macrolepidoptera of the world. Vol. V. The 
een Rhopalocera. Ed. A. Seitz. Alfred Kernen Verlag, Stuttgart. 1140 pp., 194 
pl. 

ELIOT, J. N. 1973. The higher classification of the Lycaenidae (Lepidoptera): A ten- 
tative arrangement. Bull. Brit. Mus. (Nat. Hist.) Entomol. 28:371-505 + 6 pl. 

FABRICIUS, J. C. 1782. Species insectorum. Vol. 2. 517 pp. 

Gray, P. H. H. 1962. ‘Giant’ scales on the wings of Thecla species (Lepidoptera: 
Lycaenidae). Entomologist 95:76. 











VOLUME 40, NUMBER 3 157 


HEMMING, F. 1967. The generic names of the butterflies and their type-species (Lep- 
idoptera: Rhopalocera). Bull. Brit. Mus. (Nat. Hist.) Entomol., suppl. 9, 509 pp. 
HEWITSON, W. C. 1862-1878. Illustrations of diurnal Lepidoptera. Lycaenidae. John 
van Voorst, London. 282 pp., 110 pl. 

KaFaTos, F. C. 1981. Structure, evolution and developmental expression of the silk- 
moth chorion multigene families. Am. Zool. 21:707-714. 

LaTHY, P. I. 19380. Notes on South American Lycaenidae, with descriptions of new 
species. Trans. Entomol. Soc. Lond. 78:133-137. 

NyHouT, H. F. 1978. Wing pattern formation in Lepidoptera: A model. J. Exp. Zool. 
206:119-136. 

1980a. Pattern formation on Lepidopteran wings: Determination of an eyespot. 

Devel. Biol. 80:267—274. 

1980b. Ontogeny of the color pattern on the wings of Precis coenia (Lepidop- 

tera: Nymphalidae). Devel. Biol. 80:275-288. 

1981. The color patterns of butterflies and moths. Sci. Am. November, pp. 140- 











151. 





1984. Colour pattern modification by coldshock in Lepidoptera. J. Embryol. 
Exp. Morph. 81:287-305. 

SCHATZ, E. & J. ROBER. 1885-1892. Die Familien und Gattungen der Tagfalter. In 
Exotische Schmetterlinge. Ed. O. Staudinger & E. Schatz. Lowensohn, Furth, Ba- 
varia. 284 pp., 50 pl. 

SOKAL, R. R. & F. J. ROHLF. 1969. Biometry. W. H. Freeman & Co., San Francisco. 
776 pp. 

STAUDINGER, O. 1884-1888. Exotische Tagfalter. In Exotische Schmetterlinge. Ed. O. 
Staudinger & E. Schatz. Lowensohn, Furth, Bavaria. 333 pp., 100 pl. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 158-163 


IDENTITY OF “AUTOGRAPHA” OTTOLENGUII DYAR AND 
OCCURRENCE OF AUTOGRAPHA BURAETICA 
(STAUDINGER) IN NORTH AMERICA 
(NOCTUIDAE: PLUSIINAE) 


J. D. LAFONTAINE 


Biosystematics Research Centre, Research Branch, 
Agriculture Canada, Ottawa, Ontario K1A 0C6 


ABSTRACT. Autographa ottolenguii Dyar is shown to be a typical Syngrapha with 
male genitalia similar to those of S. nyiwonis (Matsumura) and S. interrogationis (Lin- 
naeus). The type specimen is female but has wrongly associated male genitalia. A lec- 
totype is designated for the species. Autographa buraetica (Staudinger) is added to the 
North American noctuid fauna. 


Autographa ottolenguii Dyar has been considered an anomaly be- 
cause it combines Autographa-like male genitalia with other characters 
associated with Syngrapha. 

The species was originally described as Autographa arctica by Ot- 
tolengui (1902) at a time when Syngrapha was applied only to the 
small, diurnal species of Syngrapha with yellow hind wings. Ottolengui 
did, however, correctly associate the species with Autographa [now 
Syngrapha] interrogationis (Linnaeus), stating that the two species 
differ in details of wing markings and male genitalia. Dyar (1903) 
renamed the species Autographa ottolenguii because A. arctica Ot- 
tolengui is a secondary homonym of the congeneric Plusia arctica 
Moschler, now considered a synonym of Syngrapha u-aureum (Gue- 
nee). McDunnough (1944) left the species in Autographa but stated 
that it was one of two species that he had not examined. Eichlin and 
Cunningham (1978) transferred the species to Syngrapha because of 
the Syngrapha-like female genitalia and tibial spining. They treated 
it as the most primitive member of Syngrapha because of the Auto- 
grapha-like male genitalia. The species was returned to Autographa 
by Franclemont and Todd (1988), presumably because of the male 
genital characters. 

I became interested in the problem while working on the Noctuidae 
of the Beringian area. Specimens in the Canadian National Collection 
(CNC) identified as A. ottolenguii lacked the characters typical of 
Syngrapha discussed by Eichlin and Cunningham (1978). On exami- 
nation of the type specimen in the United States National Museum of 
Natural History, Washington, D.C. (USNM), it was immediately ob- 
vious that the CNC specimens were not conspecific. The overall ap- 
pearance of the type was similar to that of the circumpolar Syngrapha 
interrogationis (Linnaeus) and to that of S. nyiwonis (Matsumura 1925) 


VOLUME 40, NUMBER 3 159 





Fics. 1-4. Syngrapha and Autographa adults. 1, Syngrapha ottolenguii (Dyar), 2 
lectotype of Autographa arctica Ottolengui, Attu Island, Alaska; 2, S. ottolenguii (Dyar), 
6, Alaska; 3, Autographa buraetica (Stgr.), 9, U.S.S.R., East Siberia, Mondy, Buryatskaya, 
A.S.S.R.; 4, A. buraetica (Stgr.), 2, Palmer, Alaska. 


of the eastern Palaearctic. The Autographa-like genitalia of the type 
(Eichlin & Cunningham 1978: fig. 68) seemed inconsistent with the 
otherwise typical Syngrapha-like appearance of the specimen, and I 
began to suspect that the abdomen and genital preparation were not 
correctly associated with the adult. Three things confirmed this suspi- 
cion: first, the male genitalia are indistinguishable from those of Au- 
tographa californica (Speyer); second, a second male (Fig. 2), when 
dissected, had genitalia typical of Syngrapha (Fig. 5); third, the sex of 
the type specimen was redetermined as female, based on the brushlike 
frenulum. 


IDENTITY OF SYNGRAPHA OTTOLENGUII 


The male genitalia of Syngrapha ottolenguii (Fig. 5) confirm the 
placement of the species in Syngrapha. Within the North American 
fauna, they are most like those of S. interrogationis, but can be distin- 
guished by the characters in Table 1. Syngrapha ottolenguii is most 


TABLE 1. Comparison of male genitalia of Syngrapha spp. 


S. ottolenguii S. nyiwonis S. interrogationis 
Character (2 specimens) (3 specimens) (30 specimens) 

Basal cornutus straight, straight, absent 

(apical in vesica 1.2 mm long 1.2 mm long 

when everted) 
Apical cornutus curved, curved, straight, 

(basal in vesica 0.6 mm long 0.6 mm long 0.4 mm long 

when everted) 
Apex of valve pointed pointed blunt 
Ampulla straight, straight, recurved, 

¥% valve width % valve width %4 valve width 


Apical third of valve slightly expanded narrowed slightly expanded 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 5-8. Male Syngrapha and Autographa genitalia with aedeagus removed and 
shown at right with vesica everted. 5, Syngrapha ottolenguii (Dyar), Alaska; 6, S. nyji- 
wonis (Mats.), U.S.S.R., Magadenskaya Oblast’, Kava River; 7, Autographa buraetica 
(Stgr.), Yukon, Dawson; 8, A. pulchrina (Haw.), England. 


similar to S. nyiwonis (Fig. 6). The male genitalia of the two species 
differ only in the shape of the valve apex and in the length of the 
ampulla. The male genitalia of the three species are compared in Ta- 
ble 1. 

Adults of S. ottolenguii can be distinguished from those of S. inter- 
rogationis and S. nyiwonis by the brownish gray rather than silver- 
gray forewing ground color, and by the relatively straight transverse 
posterior line on the forewing (Figs. 1, 2). 

Ottolengui described Autographa arctica from eight specimens in 
the USNM. It is not clear from the original description that one spec- 
imen was selected as holotype. Actually, one is labeled type, the others 
are labeled co-type. To avoid confusion, I here designate the specimen 
labeled type as lectotype. It is a female in good condition except that 


VOLUME 40, NUMBER 3 161 





10 


(O 


Fics. 9-10. Female Autographa genitalia. 9, A. buraetica (Stgr.), N.W.T., Norman 
Wells; 10, A. pulchrina (Haw.), England. 


antennae and abdomen are missing. The abdomen and male genital 
preparation associated with the specimen are not from the lectotype. 
The specimen is labeled ““Type No. 6258 U.S.N.M.; 6 genit. on slide 
20 Aug. 1936 JFGC 541; Genit. slide USNM 40288; Plusia arctica 
(1902) Type Ottol.” 

The type series was nominally collected on Alter Islands, Alaska, 8 
Sept. 1880, by L. M. Turner (Ottolengui 1902). All specimens from 
mainland Alaska and Yukon attributed to this species have been re- 
identified as Autographa buraetica (Stgr.), discussed below. As a result, 
Syngrapha ottolenguii is known only from the type locality. After 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


unsuccessfully trying to locate Alter Islands, I contacted Robert Poole, 
who was able to provide critical information from the United States 
National Archives. Correspondence received by USNM in 1881 from 
L. M. Turner states that material collected in 1880 is from Attu, in the 
Aleutian Islands. Also, Turner’s handwriting in this correspondence 
and on the specimen label makes Attu look like Alter because the first 
“t’? is not crossed and the form of “‘u’ resembles “er.” 

Syngrapha ottenlenguii is known only from Attu, and may be re- 
stricted to the outer Aleutian Islands. The species likely originated in 
the eastern Palaearctic 800 km to the west where its sister species S. 
nyiwonis occurs, rather than in mainland Alaska 2400 km to the east. 

I believe this is the only noctuid known from the outer Aleutian 
Islands. 


AUTOGRAPHA BURAETICA (STAUDINGER 1892) 


Having established the identity of Syngrapha ottolenguii, I return 
to the original problem of the identity of the CNC specimens. These 
are Autographa buraetica (Staudinger), a species of the eastern Pa- 
laearctic not previously reported from North America. It can be added 
to a growing list of palaearctic species now known to occur in the 
Nearctic in Alaska and Yukon (Figs. 3, 4). Autographa buraetica is 
similar to A. pulchrina (Haworth) of the western Palaearctic but differs 
in having the thoracic tufting and forewing ground color gray-brown 
rather than bright reddish brown, in the length of the basal portion of 
the vesica (Figs. 7, 8) and in the corresponding length of the female 
ductus bursae (Figs. 9, 10). Thirteen North American specimens of A. 
buraetica in the CNC are from Palmer and Fairbanks, Alaska; Dry 
Creek, Dawson and Teslin, Yukon; Atlin in northern British Columbia; 
and Norman Wells in the western Northwest Territories. Collecting 
dates range from 27 June to 22 August. There are also two specimens 
from Alaska in USNM; these are from Palmer and Matamusaka. 

In the North American Check List (Franclemont & Todd 1983), 
Syngrapha ottolenguii should be listed after S. interrogationis and 
Autographa buraetica after A. pseudogamma (Grote). 


ACKNOWLEDGMENTS 


I thank V. S. Kononenko, Far East Science Center of the Academy of Sciences of the 
U.S.S.R., Vladivostok, and R. W. Poole, United States Dept. of Agriculture, Washington, 
D.C., for the loan of specimens; the latter also provided critical information on the type 
locality of Syngrapha ottolenguii. I also thank K. Mikkola, University of Helsinki, Fin- 
land, for assistance in the identification of Autographa buraetica; he, and P. T. Dang, 
Canadian Forestry Service, Ottawa, read and commented on a draft of the manuscript. 


VOLUME 40, NUMBER 3 163 


LITERATURE CITED 


Dyar, H. G. 1908. A list of North American Lepidoptera and key to the literature of 
this order of insects. Bull. U.S. Dep. Agric. No. 52. 723 pp. 

EICHLIN, T. D. & H. B. CUNNINGHAM. 1978. The Plusiinae (Lepidoptera: Noctuidae) 
of America north of Mexico, emphasizing genitalic and larval morphology. Tech. 
Bull. U.S. Dept. Agric. No. 1567. 122 pp. 

FRANCLEMONT, J. G. & E. L. Topp. 1983. Noctuidae. In: Hodges, R. W. et al., Check 
list of the Lepidoptera of America north of Mexico. E. W. Classey Ltd., and the 
Wedge Entomological Research Foundation, London. 284 pp. 

McCDUNNOUGH, J. 1944. Revision of the North American genera and species of the 
phalaenid subfamily Plusiinae (Lepidoptera). Mem. Sth. Calif. Acad. Sci. 2:175-232. 

OTTOLENGUI, R. 1902. Plusia and allied genera with descriptions of new species. J. 
N.Y. Entomol. Soc. 10:57-82. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 164-187 


THE LOCATION OF MONARCH BUTTERFLY 
(DANAUS PLEXIPPUS L.) OVERWINTERING 
COLONIES IN MEXICO IN RELATION TO 

TOPOGRAPHY AND CLIMATE | 


WILLIAM H. CALVERT AND LINCOLN P. BROWER 


Department of Zoology, University of Florida, 
Gainesville, Florida 32611 


ABSTRACT. Each year monarch butterflies migrate from breeding grounds in the 
United States and Canada to the Transvolcanic Belt of central Mexico. Here, within the 
montane fir forests, they initially aggregate in small groups of loose clusters scattered 
along high ridge crests. During November and December the numerous small groups 
consolidate into a few large compact aggregations and move downward into more pro- 
tected positions closer to water. Butterfly activity increases in the last half of February 
due to seasonal warming. The consolidation and compaction processes that marked the 
beginning of the season reverse, and the colonies spread out and often split into two or 
more parts. After mid-March, colony size decreases as the butterflies begin to remigrate 
northward. Several characteristics of the climate and physiography of the Transvolcanic 
Belt, including moisture, altitude, and slope exposure and inclination, are important to 
the overwintering biology of the monarch butterfly. The forests of the zone play a major 
role in satisfying the overwintering monarchs’ microclimatic requirements by moderating 
temperature extremes and conserving moisture. By colonizing this high altitude area in 
the tropics, the butterflies appear to satisfy microclimatic requirements that include 
temperatures low enough to keep activity, metabolism, and lipid expenditure to a min- 
imum, but not so cold as to cause freezing; sufficient solar input to allow thermoregulatory 
basking and consequent flight; and sources of moisture and nectar. 


Each autumn, millions of monarch butterflies (Danaus plexippus L.) 
migrate southwest or south (Urquhart & Urquhart 1978, Schmidt-Koe- 
nig 1979) from breeding grounds in eastern and central United States 
and southern Canada to overwintering sites in Mexico. Funneling 
through Texas, they cross into Mexico and encounter the southern 
extension of the Rocky Mountains, the Sierra Madre Oriental. Here 
they change their southwesterly course and follow the ranges to the 
southeast, eventually cross them, and continue to the Transvolcanic 
Belt, the volcanic mountains that extend across the southern end of 
Mexico’s Central Plateau (Altiplanicie Mexicana) between 19° and 20°N 
latitude. At a few isolated places within the high altitude coniferous 
forests, which are scattered through this belt of mountains (Fig. 1), 
monarchs spend the winter in aggregations estimated to be in the tens 
of millions (Brower et al. 1977, Calvert, in prep.). 

Monarchs migrate south in the fall to avoid winter cold and survive 
in cool, moist places where they can conserve fuel reserves in a state 
of reproductive inactivity until making the return trip north in the 
spring. Yet weather in the overwintering areas does not ideally meet 
monarch requirements. Not only do temperatures occasionally fall into 
the lethal range (Calvert et al. 1983), but also intense insolation on 


VOLUME 40, NUMBER 3 165 


clear and partly cloudy days stimulates butterfly activity to an extent 
that appears to contradict their need to conserve fuel. In an attempt 
to resolve these apparent contradictions, and to understand better why 
the monarchs choose these particular areas in Mexico, we here describe 
characteristics of the annual overwintering cycle and ecological fea- 
tures of several overwintering areas that we studied for nine seasons 
(December 1976 through spring 1985). 


PHYSIOGRAPHIC FEATURES, CLIMATE AND VEGETATION 


Volcanic cones and ranges dominate the terrain of the Transvolcanic 
Belt, which has an area of 60,000 km?, and measures approximately 
640 km across by 95 km wide (Moore 1945). To the north it is bounded 
by the high Mexican plateau, and on the south by the large Balsas 
River drainage (Rzedowski 1978). Its eastern portion averages 2200 m 
elevation with numerous peaks rising above 3600 m, including the 
highest mountains in North America south of Alaska (Goldman 1951). 
The western portion contains fewer high peaks, and declines in ele- 
vation towards the Pacific. The central area where the monarch colo- 
nies are located (Fig. 1) is drained to the north and east by the Rio 
Lerma and to the south and west by the Balsas-Mezcala river system 
(Arbingast et al. 1975). 

Classic wet-dry season weather patterns prevail through most of the 
Transvolcanic Belt. Precipitation and heavy clouding is frequent from 
May until October, especially in the mountains, but winters are dry, 
and arid conditions prevail on the interior plains (Goldman & Moore 
1945). However, winter and early spring storms occasionally occur in 
the area, and the higher elevations are subjected to high winds, heavy 
rains, snow, and ice storms (Mosina-Aleman & Garcia 1974). While 
potentially lethal to the overwintering butterflies (Calvert et al. 1983), 
these storms are also beneficial because they reduce the severity of the 
winter drought in the high-elevation overwintering areas. 

Because of the wide range of altitudes and climatic conditions, vege- 
tation within the Transvolcanic biotic province is extremely varied. 
High interior plains and valleys consist largely of grasslands intermixed 
with patches of small trees, shrubs, yuccas, agaves and cacti. On moun- 
tainous slopes, forests dominated by oaks and pines give way to firs at 
about 2750 m (Goldman 1951), but in more humid areas, the firs 
commence as low as 2400 m (Rzedowski 1978). On the highest peaks, 
firs give way to alders and other species of pine and eventually to 
grassland and tundra (Goldman 1951, Goldman & Moore 1945). As is 
true of the lower limits, vegetational transitions depend on moisture 
and exposure, and the altitudinal limits of the fir zone may be influ- 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


166 


(Dso1s a4 saiqy) [aureAo pure sui3}yed eseuleip 


Mo66 





Byyneng 


(| 


oy) \ 
oh 
qs 


a 
YW 
ENS 








Bnyjo5B4zZ) 








(esojByjes sejqy) 
1S9104 |OWBAG &! 
AWD a 
uoljB6eiBBy peyodey Vv 
UONBD07 AuojIODN ©@ 


N02 





Mo66 


eZ 


ebeuleig 





“UW OOS = [BAr9}UT m0jU0ZD 
‘(I86T) ‘uouy uO paseq st s}sa107 JouTeo ey) JO UORNIASIp pue peuljiepun ae sotuojoo JO soureN ‘Weg orueojo 


‘Aydeiso0do} 0} UOTIB[AI Ul seaIe BUTI9}UIMIZAO Ayte}4nq yoreuo 


MoOOlL 


LOS 





“000 > 





JOAIY 


IBS B] Op 
uBede}x| 





00%" 





BOBABUIEND 


sesjeg 




























ASUBI], SODIX9J] JO S}sa10j 
W jo UOReD0T “TOL 

































woose No6L 
Bonjoy oppr | Ax? sewojeg M0095 
es Y ooaz 
uy 
205, 
0 obj ep!iH ‘Po 
seupuy ues 
o6/epiy 
ep 010 13 
ONBABIB)!W woooe 
9edeyu09 O 
ousswery \ 932 
pon we NoOd 





167 


VOLUME 40, NUMBER 3 


you 0} peyersrutea1 AUOjOD _- as —- 000 Adv cI 
== aa z= OG FPN GG 
o1ajedeZ OAOIIY JO UW10}}0g — — — CLo INE 
a mE VESTS GSS 9946 
= Saas Goo ue CT 
Sait De 7s 666 9d SI 
enoulyD e11aIg ‘o1ayede7 oAoIy jo adojg = OE OT9T O8T'E TS’ AON GT — fp BnoulyD Or 
enouly) eiraig “eyourlg ey] oAo11y jo adojs OGG 00.66 O86S O10 %°d 9 ¢ enouryy) 6 
aya Se ae 6S$O 4ON GI 
enoulyD eIIeIg “eyour[g ey] oAoIIy jo odojsg ot 61 0866 O966 910 AON F G Bnouryy) 8 
enoulYyD e1leIg “eyourlg ey OAOLY o9&G KONORG WE uef TT 
SOGOTRI[IA UoouTy Jo adojsg coos SSI GREE 6FT %d 9 [ enouryy) jl, 
ag Sa EES ec case gE OLAS Oa ere eg eg ese sapere ame 
(M.L9066 “N,TTo6T) °° 
-IxoJ ‘OAPIG OP AeA Iev9U “epeiIoP{ VIPETg O119D Jo odojs BONG OSlc S41 Coo uel FZ peprlloy] 9 
(M.,9TOOT “N,8Z6T) OF 
-xoJ “eLIansy OyeUOG eeu ‘UO[ag O119D UO UOAUBO MOTIeYS ol SE FSI O000'S LT uel 1g euolad G 
(AA.,80000T “N,8So6T) 9F0 994 FT 
uvovoyoly ‘oedayuo0D 1eveu “ouRIIUIeyY O119D jo odojs 0096 0896 ZGSTS 690 AON 13 OuRITUTeITy p 
auIes JO W10}}0g SLO IeW OS 
BnoUuIyL) eIIaIg “eyour[d ey OAoIIy jo adojs OTS 0986 PESTS 69S 4H YI G enouryy) +3) 
o6&E 9L0 YH G 
enouly) PIIeIg ‘o1ayedeZ ofoIIy Jo adojs oV 03093 LIDS LEO uel CT T Bnouryy) G 
i a a a a SUV ALILGN cages Sener Pe yep mee ae rae Sie ce a aes gr ee 
(M.8T.00T “N,1F6T) 
uBdvOYOIY “eNoUTYD e1IVIS “o10}ede7 OAOIIYW Jo odojsS 098 100093 S80°E OG'T uel 1% [ ,enoury) I 
PERE eT eee EEE HEM Ba CROLL Regt Lin CLO erie ee SEs GA ME = BOR ee 
uolje00] ayeuNIxo1ddy ynuize = uoleurjoap (u1) (ey) poddeu a3eq 9suieu AuoloD ‘ON 
ule adojs apnqyy[y  9ZIS 


‘C86I-9LEI ‘OOLXeJ] Ul SATUO[OD BUTIOJUIMIGAO YOIVUOU YG JO SOIjsiiojoVIeYO [eoIshyg ‘| ATAV I, 








a 
sal 
= 
QD 
fe) 
yn eno — — = cog uefsg 
2 -UIYD PII9IG ‘o10yede7Z OAOIIY JO W0}0q IvoU adojs MOTTeYS = 19S OFEl FEOS 6FO 2d8 g enouryy) PC 
a enoulyD e1etg “eyourld ey oAoiry jo adojg~— C6 OPIS ILTS SIO ACN OF ¢ enous SG 
> enoUlyD eIIeIg “eyourld ey ofo11y jo adojg ~— 6 LT OF8Z LESS SEO AON 6 y enoury® GG 
5 enoulyD eIIIg “eyourld ey] oko11y jo adojg ~—s O6T 00S O066E S00 AON ZZ & eHouly® 1G 
= enoulyD eIIOIg “eyouryd ey] oo1ry jo adojg = SHS SST 32IS STO APN ZZ G enous) 0G 
5 enoulyD ei1IeIg “eyourld eT] oAo11y jo adojg ~— F0 FSS GSTS G60 APN ZZ [ Bnoury) 61 
eS) 9 ea ag US OSC gsc ae rg a es gece ee eg eg cee 
a (A.,98.00T “N.LP6T) 
= uBdROYOI] ‘soljnsy soy Ivau ‘saipuy ues O119D jo edo[g = 4.04 O10Z OCLs 610 %d8 saipuy ues ST 
© QI-FI Sefuofoo wio1y poatiop ‘eyourlg ey] oko11y jo Woyog 6 EZ 00FT 8263 FSO uel LT (7 AON |S) Ail 
z eyourd ey] oAorry jo adojs addq) QZ TLE ISZS GZO uel[gl gE BnouTYyD QT 
2 = Se Oro uel gt 
5 a Se GG 
a eyourld ey oAorry jo edojs addq) 6 1G 088 6S1E S80 AON 6 6 WUD) Si 
= eee ele soo uel 1z 
=e a ae sro urfg 
enoulyD eiI9Ig ‘eyouelg ey] ofo11y jo adojs 13ddQ Sg 10 00.0€ 0628 OZI AON 6 I enouryy) FI 
eae wr ee ae OOH OHS ee aarp SS a ee 
uevovoyolyy ‘oedayuor) IevaU ‘OURITUIeYY OLI9aD Jo edo[S HFG 0896 O0OCE 6IO %GAFI £OUPIIWIEITYy eT 
enoulyD eiLeIg ‘eyourlg ey] OAoiry jo adojs 19ddq — — — 60'0 AON CI g enoulyy ZI 
enoulyy) eLIVIg ‘eyourlg ey] oAOI1y jo adojs 1addq — — — cC0';0 AON CT G enouly) ital 
uoneoo] ayeullxoiddy ynulize  uodl]eUl;oap (u1) (ey) peddeur ajyeq oureu AuojoD ‘ON 
ule] adojs apnywypy  aZIs 








‘ponutjuoy =‘T ATAV 


168 


169 


VOLUME 40, NUMBER 3 


‘sUOTLIAAp PAepue}s puke SUBUT OU} ayndwi0o 0} uasoyo sem (potsed ayqeis ey) Jo a[ppru ayy) Arenaqay [ 07 ysasopo yey 
(6L6I [Be 19 Wea[eD) 


(M.8T000T 
‘NU ZEo6T) UBoVOYoT ‘odurwoO 1eveU ‘TWBATYD) OLIID) JO adojs 


(A\,00000T ‘N,00.6T) Odue0O e98u 
‘oreuvdurey [A PIII ‘solauor) soy oAOIIy fo adoj[s 


(M,STOOT ‘N,GS6T) UROBOTSTIN ‘odureoQ iveu ‘orreu 
-edwiey) [A PLIaIS ‘epusloey eT ofoi1y jo siayeMpeoyy 


eyourld ey oAorry Jo edojs 


(AA, ST00T 
‘N 68061) PUNY SI sureU [BOO] ‘UBOBOYOTN, “9PUBID) 


oyouey eile ‘soutpndeD so] C119) J AS uleyUNOY 
eyourlg ey] OAOITY JO W10}0q IvON] 





woTyZI0] ayeuNIxo1ddy 


o8G all 

TOSS 1GGoGS 
oFSI1 0SoLT 
o6GG O1oh%S 
ESGS OGoLT 
OST OFoSS 
68/1861 
= 00.0 
OSI 0001S 


ynulize 
ule 


uoljeUuTpoep 


adojs 


sil v6 0 
SsTs S80 
8608 020 
SLITS PEs 
CSTS sO 
1466 LVI 
Ors = 

000s 19% 
EOS — 

sie, 09°0 


(ud) (ey) poeddeur a3eq 


apnynty —_9ZIS 


} ‘Uayxe) SEM JUSUIAINSeAU 9UO UPY) P1OUL 9194 MM « 
vare BIC AUS [PUIBIIO OY] , 


(6LOI [Be 28 Woealeo) Bele BUTWURS) 9}IS [PUIBLIO OY L « 
(6LOL JB J2 WeAaleO) Bole Bog PIS [Bulsiio oY z 
(LLOI [Be 12 Jamoig) vole eydiy ays [eulsi0 94] | 


[eAsioqul 





HearyO 


o1reueduie) 


T oyorolg 
enoulyy) 


eunoy 
L enoury) 


auieu AUOlOD 


aouspyuoo 
iejnsuy 10 GS 
cUBOIN\ 


O& 
66 


86 
LG 








Eee See ee ee a SS ae ae eee 


‘ponuljuor) 


‘T W1dV.L 


170 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


enced by the presence and extent of the summer fog belt (review in 
Brower 1985). 


HISTORY OF THE OCCURRENCE OF OVERWINTERING COLONIES 


Residents of settlements located near overwintering colonies in the 
Sierra Chincua, Michoacan, claim that monarchs have always come to 
these areas. Although butterfly motifs occur widely in Precolumbian 
art and mythology (Brewer 1983), especially in the Teotihuacan cul- 
ture (Castellanos 1983, de la Maza 1976), we have not found mention 
of monarch colonies in the literature prior to Urquhart’s original report 
(1976). Lack of Precolumbian records of the conspicuous overwintering 
phenomenon may be due in part to its location near the boundaries of 
the Tarascan and Aztec empires (West 1964), which may well have 
been a dangerous no man’s land. Possible folk-knowledge of monarchs 
may be expressed in the frequent use of the local Spanish name for 
butterfly, “paloma,” in topographic features both in the overwintering 
areas and in areas through which they migrate. The migratory phe- 
nomenon has also found its way into the language of the Mazahua, a 
group of Indians living in the migratory corridor in the village of 
Santiago north of Villa Victoria, in the state of Mexico. Their word, 
“Seperito,” translates as “‘butterfly that passes in October and Novem- 
ber’ (Kiemele Muro 1975). Historically, monarchs may have been im- 
portant to the Mazahua as a supplemental food source. Collecting them 
at temporary roosts during the fall migration, Mazahuas still eat mon- 
archs after removing the wings and frying them on flat ceramic pans 
(comals). However, the practice is now apparently limited to occasional 
performances for tourists (Yamaguchi 1980). 


METHODS 


We here depart from procedures in our previous report (Calvert et 
al. 1979) and name the colonies according to the mountain peak or 
range on which they are located. In cases where two or more colonies 
occupied the same peak or range in the same year, a number follows 
the name and signifies a specific colony location with the peak or range. 
Site Alpha, originally described in Brower et al. (1977), was located in 
the Sierra Chincua and is now called the Chincua overwintering area. 
Place names, geographical features and coordinates (Table 1) were 
determined from the Mexican CETENAL map series (Anon. 1976a). 

Between December 1976 and March 1982 we spent a total of 19 
months at various colonies in the Transvolcanic Belt including 94 days 
during 1978-79 at Chincua 4 in the state of Michoacan. During this 
time we located 30 colonies on 5 mountain massifs and mapped each 
using a Suunto sighting compass and a 100 m surveying tape. Colony 


VOLUME 40, NUMBER 3 1 7Aal 


area was computed using a Hewlett-Packard double meridian distance 
program or an Apple II graphics program. Colony boundaries were 
marked with date-coded colored tape to monitor changes in positions. 
We recorded the declination of the mountain slope at the position of 
the colony and the “facing azimuth,” that is, the direction of the down- 
slope line perpendicular to the contour at the colony center. To deter- 
mine significance and angular confidence interval of the average facing 
azimuth, we applied circular statistics (Batschelet 1972) to azimuths, 
corrected for magnetic declination, approximately 8.5° east (Anon. 
1976a). Circular statistics were also used to derive the angular confi- 
dence interval for the average slope declination. 

During 1978-79, daily temperatures were monitored continuously 
from 19 January—24 March with recording hygrothermographs (Brow- 
er & Calvert 1985) at two locations, one in the forested center of 
Chincua 4 (Colony #10, Table 1) and the other in a nearby clearing. 
Forest and understory plants were identified at the University of Texas 
Lundell Herbarium, or by reference to Sanchez (1979). 


RESULTS AND DISCUSSION 
Location of Colonies 


The 30 colonies we found were located in the high-altitude moun- 
tainous terrain of the Mexican Transvolcanic Belt between 19°10’ and 
20°00'N latitude and 99°55’ and 100°40’W longitude (Table 1), a rec- 
tangle of about 7000 km? (Fig. 1). Evidence of other colonies, indicated 
by the presence of detached wings and body parts spread over areas 
up to 0.25 ha, occurred as far east as 99°52’ near the volcano Nevado 
de Toluca. (A small colony indicated as Los Palomas in Fig. 1 was 
discovered here in November 1984.) Small overwintering aggregations 
confined to one or a few trees have been reported east of Mexico City 
on the western slopes of the volcanoes Popocatepetl and Ixtaccihuatl 
(98°45'W;; J. de la Maza, pers. comm.) and south of Mexico City in the 
vicinity of Tres Marias (99°10’W; J. Mausan, pers. comm.). These small 
aggregations appear to be outlying groups that do not form every year, 
and no mass movement of migrant butterflies into these areas has been 
observed or reported. 

Several locations outside the Transvolcanic Belt, and in its eastern 
extreme, appear to have habitat characteristics and the requisite alti- 
tude to be suitable for monarch colonies. Monarchs occasionally are 
seen in migration towards areas removed from known colonies. For 
example, in October 1980, we saw large numbers migrating SE along 
the escarpment above Orizaba, Veracruz, apparently headed to the 
Sierra de Juarez, Oaxaca. In 1977 another group was observed far from 
known overwintering areas and migratory pathways flying ESE near 


iy JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Candelaria Loxicha, Oaxaca (de la Maza et al. 1977). Accordingly, we 
searched in several mountain ranges outside the known overwintering 
areas, including the western slopes of the Cerro Pena Nevada in the 
Sierra Madre Oriental, state of Nuevo Leon, in December 1977, and 
the western slopes of the Cofre de Perote and the Pico de Orizaba, in 
the states of Puebla and Veracruz in February 1980. In February 1982, 
our group, including Javier de la Maza of the Mexican Department of 
Wildlife, investigated several areas in the Sierra de Juarez and the 
Sierra Madre del Sur, all in the state of Oaxaca. De la Maza returned 
to these areas in March and early April of the same year and explored 
further south in the Sierra de Chiapas, eventually reaching the moun- 
tainous border of Guatemala. No colonies or evidence of colonies (such 
as large numbers of dismembered wings) were found. 

Unless further data are forthcoming, we conclude that the principal 
overwintering colonies of the eastern population of the North Ameri- 
can monarch (Fig. 1: Pelon, Chivati, Picacho, Campanario, and Chin- 
cua) occur on a few isolated mountain ranges confined to a remarkably 
small area of approximately 800 km? between 19°20’ and 19°45’N lat- 
itude and 100°10’ and 100°20’W longitude (at this longitude 1° of lat- 
itude = 111 km and 1° of longitude = 104 km). Scattered aggregations, 
such as colonies Altamirano, Herrada and San Andres (Table 1) are 
found outside this area, but these are always small, do not form every 
year, and break up earlier than those in the central area. 


Colony Vegetation 


Most places in the Transvolcanic Belt between 2500 and 3500 m 
elevation are dominated by the “oyamel’’ fir (Abies religiosa H.B.K.), 
which occurs on all terrain except rocky outcroppings and areas of cold 
air drainage (llanos). This is the major species upon which monarchs 
form their overwintering roosts. Other trees in the community include 
Pinus pseudostrobus Lindl. (referred to as Pinus ayacahuite in Brower 
et al. 1977), Cupressus lindleii Klotzsch, several species of Quercus, 
and Buddleia cordata H.B.K. (the last species is confined to moist 
canyon bottoms). Cupressus lindleii is found in dense stands at lower 
elevations within Abies religiosa forests, while Pinus pseudostrobus 
and Quercus spp. are more scattered and appear as occasional individ- 
uals within the Abies forest. Monarchs occasionally roost on all of these 
tree species but, since they mostly locate their colonies in oyamel- 
dominated forests, they are usually found on the oyamels. Moreover, 
when used as roosting trees, the broadleafed angiosperms are generally 
not covered as densely as are the conifers (Brower et al. 1977). 

The most conspicuous components of the forest understory include 
tall (4 m) and medium (2 m) woody shrubs, dominated by composites, 


VOLUME 40, NUMBER 3 Eis 


the most important of which are Senecio anquilifolius D.C., S. barba- 
Johannis D.C., Eupatorium mairetianum D.C. and E. patzcuarense 
H.B.K. Important noncomposites in this portion of the understory are 
Cestrum anagyris Dun., Salvia elegans Vahl. and S. cardinalis H.B.K. 
Ground cover is dominated by Alchemilla procumbens Rose and one 
or more species of mosses in the genera Thuidium and Mnium. Local 
clearings contain Senecio stoechadiformis D.C., S. tolucanus D.C., S. 
prenanthoides A. Rich., S. sanquisorbae D.C., Eupatorium sp. and 
Baccharis conferta H.B.K. All of the species above, except Alchemilla 
procumbens, the Salvia, and the mosses, serve as monarch nectar sources 
at some time in the overwintering period. They also serve as substrates 
for drinking dew, especially in November and December. 

Human disturbance is common in these forests. Lumbering without 
clearcutting has been practiced for many years in all overwintering 
areas, and many forest fires have occurred, attributable to lightning, 
human carelessness and, in some areas, to the practice of renewing and 
extending pastures. Tree densities range from over 1000/ha in young 
stands to about 150/ha in severely thinned forests (Calvert et al. 1982). 


Colony Formation and Movement 


In the Sierra Chincua, we observed colonies forming only within a 
narrow time range, from 2-9 November. Local foresters near the Sierra 
E] Campanario report that butterfly arrival may vary among years by 
as much as two weeks and that they have arrived as early as the third 
week of October (P. Silva, pers. comm.). Initially, the monarchs ag- 
gregate in numerous small nuclear groups along or just below mountain 
ridges in dense foliage on the tops and sides of the trees. In the Sierra 
Chincua, dozens of these small groups form along the major NW-SE 
ridge above the Arroyo La Plancha and its two northwestern extensions 
on both sides of the Arroyo El Zapatero. 

This initial phase of colony formation is characterized by much 
movement and intense flight activity. We hypothesize that the pres- 
ence of small groups along high ridges serves as a visual cue to attract 
more migrants to the roosts. Apparent signalling during this early part 
of the overwintering season may also occur through an additional and 
most remarkable behavior in which the butterflies form large towering 
spirals. We have seen these columns of soaring butterflies in early 
November extending vertically at least to the limits of 10x binoculars 
(300 m) above ridges and clearings near colonies which were in the 
process of formation. 

The many small nuclear groups may move about and reform at other 
locations or coalesce with others, and by early November they are 
scattered along the ridges just below the crests for distances up to 3-4 


174 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fic. 2. Trees coated with thick layer of rime and ice on a ridge above Chincua 6 
(Colony 24, Table 1), 24 January 1981. The colony is located just outside photograph to 
the lower right. Exposed trees along the ridge are those on which first colony formation 
frequently takes place. 


km. Movement continues into December and gradually the many 
smaller groups coalesce into a few (1-5) larger ones located further 
downslope, but always less than 200 m below the ridge crests. Fall 
migrants continue arriving during November and early December. 

Consolidation into these larger groups continues through December 
with some colonies increasing in size at the expense of others. For 
example, during 1978/79, six groups were in the Chincua area on or 
before 6 December (Table 1). By early January, four had almost dis- 
appeared, and at least one of the two remaining ones had grown. 
During 1980-81, all five colonies at Chincua mapped in November 
had moved by 11 January from their positions on the upper slopes of 
Arroyo La Plancha to form a large (2.51 ha) colony lower down in the 
canyon (Table 1). 

A major advantage of the movement of colonies from positions on 
high ridges to lower protected areas is the avoidance of the high allti- 
tude southwest winds and occasional storms that occur during the over- 
wintering season (Mosino Aleman & Garcia 1973). This was dramati- 
cally illustrated in January 1981, when moisture-laden, gale-force winds 
covered the vegetation on an exposed ridge, where all of the groups 
had initially formed, with heavy ice and rime (Fig. 2). Note that the 


VOLUME 40, NUMBER 3 175 


rime and ice did not form on trees in the protected valley below the 
ridge where the butterflies had established their colony. But even such 
sheltered butterflies can be dislodged by the tens of thousands from 
their roosts by high winds, rain, snow, or hail (Calvert et al. 1988). 
Once on the ground, they are subject to increased risk of mortality 
from freezing (Calvert & Brower 1981, Calvert & Cohen 1983) as well 
as mouse predation (Brower et al. 1985). 

During the first phase of colony formation in November and De- 
cember, the butterflies usually occupy only the outer periphery of tree 
branches, and rarely settle on trunks. However, by early to mid-Jan- 
uary, they pack onto trunks and branches of the mid-sections of trees, 
avoid the upper quarters of crowns, and usually also avoid the lower 
branches. Packing into the interior branches and onto the trunks results 
in decreasing the total area occupied by the colony (Fig. 3a-c). On 
steeper slopes and for a short while after a colony has moved and 
resettled, clustering may occur on the lower branches as well. Cluster- 
ing on trunks occurs in two ways. During normal weather, the butter- 
flies form trunk and bough clusters in the same manner by flying up 
to a group of butterflies and landing. Following storms, the many 
thousands of dislodged individuals crawl up from the ground onto any 
vertical surface until they encounter butterflies or other obstacles. In 
this way they form the spectacular clusters which often extend nearly 
the full length of the trunk (Fig. 4). 


Colony Location and Movement in Relation to 
Moisture Requirements 


Overwintering in the high mountainous forests of the Transvolcanic 
Belt positions the colonies in a wetter habitat than that prevailing over 
much of Mexico during the winter. These conditions are due to a 
pattern of moist subtropical air masses that move east from the Pacific 
in late winter and early spring. When these air masses encounter the 
higher mountains west of the Continental Divide, adiabatic cooling re- 
sults in cloud formation and precipitation, and a more even distribution 
of yearly precipitation than at lower elevations (Mosino Aleman & 
Garcia 1974, Anon. 1978: Landsat photo). 

In years with ample rainfall, consolidation and movement cease by 
January. The colonies are typically located in protected habitats in a 
shallow, moist canyon along or near the headwater of a stream, where 
they remain until late February or March. Some colonies persist on 
mountain slopes unassociated with depressions, but almost without ex- 
ception a source of water is located within less than 1 km. 

Notwithstanding the wetter conditions at higher elevations where 
the colonies are located, long spells of dry weather do occur. During 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 








NOV. 15, 1978 DEC. 13,1978 
AREA 2.5\ha AREA 3.29ha 
A {oO m 100 m 
-———4 4 


JAN. 15,1979 
Cc AREA 2.25 ha 
100 m 
jet 










































MAR. 22,1979 
AREA 2.72 ha 





FEB. 9,1979 
D AREA 2.25ha 
100 m 


/ MAR, 3, 1979 
n/a ARIE Amc omina 
100 m 

Passat ssee 






100 m 
-————_H 








Fic. 3. Seasonal movement and size change of colony Chincua 4 from shortly after 
its formation in fall 1978 to its breakup in spring 1979. On 22 March a third fragment 
of the colony was 1 km downstream and is not shown in F. Contour interval = 20 m. 


these times, tens of thousands of butterflies fly out of the colonies to 
drink in sunny areas along streams, moist patches or at reservoirs. 
Because of the distance of the colony from water, the butterflies must 
expend significant amounts of lipid (Chaplin & Wells 1982) in their 
watering flights. During exceptionally dry years, entire colonies may 
move downward and reform along or near water. For example, during 
1979-80, when no rain occurred in the Sierra Chincua from 15 De- 
cember to 9 February, three colonies located on the dry upper slopes 
of the Arroyo La Plancha moved about 0.5 km downslope (300 m 
elevation) during three weeks and reconsolidated into a single colony 
at the juncture of two streams (Colony 17, Table 1). 


VOLUME 40, NUMBER 3 Ler F 


In addition to these occasional large-scale movements down to mois- 
ter and more sheltered areas, colonies move incrementally or “creep”’ 
downslope during overwintering. This is explained as follows: On sun- 
ny days the thousands of butterflies that fly out of the colony return 
later the same day and reform their clusters on those portions of the 
lower colony periphery exposed to the sun. During winter afternoons, 
when most return, this is the southwest side. Cluster reformation occurs 
on the lower side of the colony because this part is usually closest to 
the water source. The net effect is a slow movement of the colony 
downslope, or down canyon, depending on where the parent colony is 
located. During periods of warm weather, when greater numbers travel 
out to water each day, this downslope creep can be rapid. 

The relation of the butterflies’ return route from water to the direc- 
tion of colony movement was made clear in one notable exception to 
the above pattern. On 14 February 1978, a small, new segment of 
Colony 4 (Table 1) was located upslope 25 m northeast of the previ- 
ously mapped colony. Instead of returning to the colony directly from 
the water source located to the south at the base of the mountain, some 
butterflies circled around the mountain and returned over the top from 
the northeast. As they approached the old colony, these butterflies 
reformed clusters on trees upslope from the old colony, and, as in the 
other instances, on branches exposed to the afternoon sun. Thus, the 
path of return from water seems to be the principal factor determining 
the direction of the incremental type of colony movement. 


Colony Breakup 


Due to seasonal warming in late February and March (Fig. 5), but- 
terfly activity increases, and the colonies begin to reverse the consoli- 
dation process as larger and larger numbers fly out to water and nectar 
sources. The increasing daily efflux results in an acceleration of down- 
slope or down-canyon movement. Often the colony splits into two or 
more parts as the butterflies returning from daily activities reform 
clusters nearer water or nectar (Fig. 83d-f). As occurs when they first 
arrive in November, new clusters form on the periphery of branches, 
while interior branches and trunks are largely avoided, resulting in a 
lower density but an increase in the total area occupied by the colony 
(Fig. 3e-f). After mid-March, colony size decreases as the butterflies 
start their remigration northward. 


The Formation and Breakup of a Colony—a Case Study 


The most detailed studies of the overwintering butterflies have been 
conducted near Site Alpha (Brower et al. 1977), located in what we 
now call the Chincua overwintering area. The Chincua Area is located 
at 19°41'N and 100°17-18'W in the Sierra Chincua 5-7 km NNW of 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 4. Packing of monarch butterflies on trunks of the oyamel fir (Abies religiosa) 
is especially noticeable after winter storms when butterflies dislodged from their clusters 
by storm action crawl up the trunks. 





VOLUME 40, NUMBER 3 179 


TEMPERATURES (°C) 


Center of Colony 
Adjacent Clearing - - - 









yp 
Vitee 
Lila 
Yl 
B22 = 
Wj 
Ya 


January February March 


DATE - 1979 


Fic. 5. Daily minimum and maximum temperatures as measured by one thermo- 
graph beneath the forest canopy in the center of a Chincua colony (Colony 10, Table 1) 
from 19 January—24 March 1979 (dashed lines), and another in a nearby clearing, from 
19 January—28 February (solid lines). Horizontal dashed line is at 15.5°C, the approximate 
temperature at which monarchs are able to raise their thoracic muscles to flight temper- 
ature by shivering. 


the town of Angangueo, Michoacan (Anon. 1976a: topographic map). 
The terrain is dominated by a SE-NW ridge of the Sierra Chincua, 
which drops from 3300 m to 2600 m elevation in its 6.8 km length, 
and is centered on a local landmark known as the Mojonera Alta (“high 
dividing line,’ Santamaria 1974). To the south, the ridge falls off steep- 
ly into the deep canyon of Arroyo La Plancha (also known locally as 
the Arroyo Hondo). The north slope of the Sierra Chincua is dissected 
by canyons that drain northwesterly. At its western end, the ridge 
divides forming the eastern and western sides of the westernmost of 
these canyons, Arroyo El Zapatero. Here butterfly colonies formed in 
seven of the past ten years, from 1976/1977 through 1985/1986. 

We monitored the formation, consolidation, and breakup of one of 
these colonies (colony 10, Table 1) during the 1978-79 overwintering 
season (Fig. 3). When located on 15 November it was on the SW- 
facing slope (230°) of the head waters of Arroyo El] Zapatero, approx- 
imately 140 m from the ridge crest and 180 m from the canyon bottom 
(Fig. 3a). At this time, the colony was still in the aggregation phase, 
and occupied 2.51 ha. By 18 December (Fig. 3b), it had moved down- 
slope approximately 70 m and occupied 3.29 ha, the largest area it 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


would attain. By 15 January (Fig. 3c), it had moved 110 m downward 
and westward. Due to increased packing on trunks and the interior 
parts of branches, it had consolidated to 2.25 ha. On 9 February (Fig. 
3d) its size had not changed, but it had moved slightly westward (20 
m) with its lower and upper boundaries remaining fixed. We mapped 
the colony again on 8 March (Fig. 3e), by which time the butterflies 
were very active. The colony had split into two parts and had spread 
out along Arroyo El Zapatero. It now occupied a total of 2.72 ha and 
was located 140 m west of its February position. We mapped the 
colony the last time on 22 March. By then it had spread out over 1 km 
of the canyon, occupied 3.2 ha, and consisted of two adjacent groups 
(Fig. 3f) and a third group farther downstream. On 15 April, walking 
the entire length of Arroyo El Zapatero, we encountered only one live 
butterfly where less than a month before there had been tens of mil- 
lions. 


Colony Location in Relation to Slope Exposure 


The distribution of facing azimuths of the colonies is shown in the 
circular diagram (Fig. 6). The mean vector was 230°18’ with a 95% 
angular confidence interval of +28°. Sixty-seven percent of the colonies 
formed on slopes in the SW quadrant, which shows a clear preference 
of the butterflies for this quadrant (r = 0.56, P < 0.001). Of the nine 
colonies that formed on slopes facing other directions, two were located 
just to the east, and in the lee of, a SW-facing ridge (Colonies 1 and 
2, Table 1), five were located in canyons that drained to the west 
(Colonies 3, 5, 25, 27 and 28) and the remaining two formed on slopes 
facing SSE, one very close to the SW quadrant (Colonies 22 and 30). 
One colony formed initially on a SSE slope and later moved around to 
a slope in the SW quadrant (Colony 8). 

During the winter in mountainous areas, southern slopes receive 
more insolation and dry faster than northern ones. Moreover, SW slopes 
are heated more than SE ones because, when the sun strikes the eastern 
slope in the morning, much of its energy is spent in evaporating water 
that precipitated as dew during the night. In contrast, the afternoon 
sun strikes relatively dry ground, so most of its energy is spent in 
heating the surface (Geiger 1950). In spite of a strong need for the 
butterflies to stay cool, conserve fuel, and avoid desiccation, they ap- 
pear to choose the hottest and driest slopes available. This apparent 
contradiction needs explanation. 

We hypothesize that their location on SW-facing slopes is a com- 
promise that satisfies several requirements. During mid-winter days, 
air temperatures within the shade of the oyamel forests rarely become 
warm enough for monarchs to fly spontaneously. The minimum am- 


VOLUME 40, NUMBER 3 18] 





Fic. 6. Direction of the slopes (faing azimuths) on which 27 monarch butterfly col- 
onies were located in Mexico’s Transvolcanic Belt, 1976-82. Shaded area is angular 
confidence interval. 


bient temperature at which monarchs can shiver and then fly is be- 
tween 12.7 and 16°C (Kammer 1970, Masters 1985). Air temperatures 
of this magnitude were not reached in shaded portions of the 1978-79 
Chincua 4 colony until March 9 (Fig. 5). Even though temperatures 
are below flight threshold within the colony during most of the over- 
wintering season, direct solar radiation usually reaches the butterflies 
as sun flecks at some time during clear or partly cloudy days. This 
allows them to warm to flight threshold by basking so they can regain 
their positions in the trees after being knocked down by storms, and 
also insures that they can fly out to water and nectar on clear days. In 
addition, their basking posture readily displays their aposematic col- 
oration and helps to deter bird predators, which killed an average of 
15,000 butterflies/day during 1978-79 (Brower & Calvert 1985). This 
same study showed that predation is inversely related to temperature 
and suggests that the birds would have an even greater advantage on 
the colder, north-facing slopes. Although a position on the north-facing 
slope would be better for maintaining water balance and conserving 
lipids, the colder temperatures would result in greater inactivity so that 
the butterflies would be less able to redress an unfavorable water bal- 
ance, reestablish colony integrity after storms, avoid predators, and 
obtain nectar. 


The Importance of Slope Inclination 


Surface heating of mountainous areas depends on the amount of 
radiation striking slopes. While several factors including season, time 


182 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


of day, latitude, and the facing direction of slope affect radiation, the 
degree of slope is critical. Schubert (in Geiger 1950) showed that during 
winter, a south slope inclined at 23.5° receives 9% more heat than a 
horizontal one. The inclination of the slopes on which the butterfly 
colonies are situated averaged 25°22’ with a 95% angular confidence 
interval of 1° (Table 1). Butterflies roosting here would receive more 
solar energy to warm them, and their ability to fly would thus be 
enhanced. 

Moreover, surface heating due to solar radiation striking near the 
perpendicular to the mountainous slopes occupied by the colonies is 
maximal on about 1 March. This coincides well with the date that the 
butterflies begin to accelerate flight activity and mating, and move 
down the canyons in preparation for their remigration to the north. 
The reason for this early March maximum is as follows: At the latitude 
of the overwintering colonies (ca. 19.5°N), the date when the sun’s rays 
are exactly perpendicular to the average colony slope (25°22' = ap- 
proximately 25.4°) is 1 March. At the spring equinox, the angle of 
incidence of the sun at the latitude of the colonies is 95.5°. This value 
is obtained by adding the average slope inclination (25.4°) to the angle 
of incidence of the sun’s rays to the horizontal [25.4° + (90° — 19.5)] = 
95.9°. Therefore, at solar noon on 22 March, the sun is 5.9° beyond the 
perpendicular to the average colony slope. Since the earth precesses at 
a rate of 0.2575°/day, it takes about 23 days (5.9°/0.2575°/day = 22.9 
days) to reach this position. Therefore, the sun’s rays were perpendic- 
ular to the colony slopes about 1 March. 


Why Do Monarchs Migrate to Mexico’s Transvolcanic Belt? 


Evidence that monarchs are not sufficiently cold-tolerant to survive 
winter conditions in their northern breeding areas derives from several 
sources. Urquhart (1960) found that 77% of monarch pupae subjected 
to freezing temperatures associated with a cold front perished. Calvert 
and Brower (1981) showed that 60% of monarchs wetted by naturally 
occurring dewfall in a clearing near a Sierra Chincua colony suffered 
flight impairment or death after one night’s exposure to temperatures 
averaging only —1.7°C. Moreover, during January, 1981 within a Sierra 
Chincua colony (Colony 24, Table 1) inclement weather with low tem- 
peratures reaching —5°C resulted in the death of an estimated 2.5 
million butterflies (Calvert et al. 1983). Anderson (in Brower, in press) 
found in a controlled freezing chamber that the temperature at which 
50% of the dry and wet monarchs freeze from a Chincua colony is 
—7.8°C and —4.4°C, respectively. Calvert et al. (in press) found in 
the same area that the temperature at which 50% of monarchs wetted 
by nighttime dewfall in the open forest freeze or are injured to the 


VOLUME 40, NUMBER 3 183 


point where they could not fly normally is —3.1°C. Thus, temperatures 
slightly below freezing kill or incapacitate monarch butterflies, neces- 
sitating their migration from their northern breeding grounds. An ad- 
ditional reason for migrating southward is the senescence of their larval 
food plants. However, if the monarchs could survive the cold, they 
could, presumably, overwinter in the north as do other adult butterflies 
such as the mourning cloak (Nymphalis antiopa L.). 

The monarchs’ need to avoid lethally cold temperatures must be 
balanced against their need for moderate cold to avoid too rapid de- 
pletion of their lipid reserves (Tuskes & Brower 1978, Chaplin & Wells 
1982, Brower 1985, Walford & Brower, in prep.). Long-term minimum 
temperature records (10-29 years) for November through March at 
seven meteorological stations above 2500 m located near the overwin- 
tering colonies averaged 2.3°C (Anon. 1976b). The average minimum, 
average maximum and mean temperatures for a 40-day period (19 
January—28 February 1979) in a clearing 20 m outside a Sierra Chincua 
colony (24, Table 10) were 2.9°, 14.9°, and 8.9°C (Fig. 5; also Brower 
& Calvert 1985). The overwintering areas are thus located at an altitude 
and latitude that normally provide a cool environment with relatively 
stable minima at or just above freezing. 

The dense forests of the Transvolcanic Belt also play an important 
role in moderating the climate by reducing daily temperature ex- 
tremes. Thus, inside the center of the colony for the same 40-day 
period described above, the average minimum temperature was higher 
(4.2°) while the average maximum and mean temperatures (11.0° and 
7.7°) were lower than the corresponding averages in the nearby clear- 
ing (compare Fig. 5). The moderating effect of the forest cover be- 
comes crucially important for monarch survival when temperatures 
plummet to the killing threshold (Calvert et al. 1983). 

Latitudinally, the monarch’s choice of colony location may be lim- 
ited by two physical barriers. Immediately south of the Transvolcanic 
Belt lies the Balsas River depression (Fig. 1). Here the hot, dry climate 
would undoubtedly break their reproductive diapause (Baker & Her- 
man 1976), and cause them to begin ovipositing in areas already oc- 
cupied by indigenous milkweed butterflies, Danaus gilippus L. and D. 
eresimus Cramer, as well as nonmigratory monarchs (Calvert, unpubl. 
obs., Brower 1985). To the north lie the dry plains of the Central 
Plateau where in winter the butterflies would be subjected to colder 
and drier conditions brought about by the influence of advective air 
masses moving down from the north (Zepeda 1941, Mosino Aleman & 
Garcia 1973). In this thinly forested region they would also not have 
the protective cover and microclimatic requisites provided by the dense 
forests of the Transvolcanic Belt. Suitable habitats may exist further 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


south in Mexico and Guatemala, but areas above 3000 m are more 
limited in extent and lack the extensive fir forests found in the Trans- 
volcanic Belt (Calvert & de la Maza, unpubl. obs., Anon. 1981: distri- 
bution of oyamel forests). 

The conflicting requirements to avoid freezing temperatures and yet 
remain relatively cool, and to avoid excessive moisture that raises the 
temperature at which freezing injury occurs, and yet to remain rela- 
tively moist to avoid desiccation are closely approximated in the oya- 
mel forest ecosystem at the geographic location between 19 and 20°N 
in Mexico’s Transvolcanic Belt. 

The monarchs’ return to the north is also bounded by several con- 
straints. It must not precede the flush of spring milkweeds and the 
recession of lethally cold, late winter temperatures. And finally, it must 
allow for the first spring generation, largely produced in the Gulf 
Coastal States (Brower et al., in prep.) to mature before the advent of 
lethally warm summer temperatures in the southern United States 
(Malcolm et al. 1987). 


(CONCLUSIONS 


Monarch migration to and colonization of high altitude tropical areas 
in Mexico appear to be an adaptive response to several temperature- 
and water-related needs. Flight capacity is necessary to regain position 
in clusters following storms, to avoid predators, to locate nectar sources, 
and, especially, to obtain water. Direct and indirect solar radiation 
augmented by a SW exposure and favorable slope inclination allow the 
butterflies to warm sufficiently to fly when temperatures within the 
forest are below flight threshold. However, warming and flying in the 
course of daily activities increases their utilization of precious lipid 
reserves. The generally cool climate with sufficiently intense sunshine 
to allow flight appears to satisfy both their need to conserve energy 
and fly out of the colony when necessary. The location of the mountains 
is far enough south to minimize the impact of most cold air masses, 
which dominate winter weather in the north. Nighttime minima, which 
seldom drop more than a few degrees below freezing except during 
occasional extreme cold periods, reduce the likelihood of death by 
freezing. These often conflicting requirements are approximately sat- 
isfied by the unique climatic and physiographic features of Mexico’s 
Transvolcanic Belt. 


ACKNOWLEDGMENTS 
We thank Gov. Cuauhtemoc Cardenas and Juan Jose Reyes R. of the Department of 


Forestry and Wildlife for permission to study the sites. Mayor Jose Martinez and the late 
Mayor Manuel Arriaga, Secretary Pablo Pina, and many inhabitants of Angangueo, Mi- 


VOLUME 40, NUMBER 3 185 


choacan, provided help during our stays in Mexico. We also thank the late Col. Timoteo 
Mondragon, his son, Mauricio Mondragon, and members of the Ejido Cerro Prieto for 
permission to set up our research camps. Billie Turner and Jackie Poole of the Lundell 
Herbarium, University of Texas at Austin, helped in plant identification. Amando Manon 
B. of the Mexican Meteorological Service generously provided weather records. Peter 
Phillips, Susan Kress, Margaret Sheppard, Paul Spitzer, Gary Mathews, John Smiley, 
Lynne McAnelly and James Mallet helped collect data. Javier de la Maza, Fred Morrison, 
Diane Mahan, Diana Watts, John Christian and Enrique Fuentes assisted in the search 
for colonies. Linda Liscomb, Betty Olds, Robin Knowlton, Patra Cianciolo, and Margaret 
and Richard Barthelemy maintained healthy and pleasant camps during many long days 
of research. Evodio de Jesus and his late father assisted in camp maintenance. We are 
especially indebted to Willow Zuchowski and Richard Lindley who helped in all stages 
of the research. Elizabeth O’Berry, Ed Klostermeyer, Linda Fink, and Alan Masters made 
suggestions that improved the manuscript. 

This study was carried out as an adjunct to research supported by U.S. National Science 
Foundation grants to Amherst College and to the University of Florida, with L. P. Brower 
as principal investigator, and by World Wildlife Fund project 1958/32981958 with Wil- 
liam H. Calvert and Javier de la Maza as coprincipal investigators. We also thank the 
Center for Field Research of Watertown, Massachusetts, and our Earthwatch collabo- 
rators. 


LITERATURE CITED 


ANON. 1978. Landsat Photo Mexico #1200-16341. Tecozautla, 8 February 1973. N20- 
14.W099-31. U.S. Dept. Interior, Geological Survey. 

ANON. 1976a. Detenal Carta Topografica, Mexico y Michoacan. Angangueo E14A26, 
Escala 1:50,000. Cetenal, San Antonio de Abad 124, Mexico 8, D.F. 

ANON. 1976b. Normales Climatologicas. Direccion General de Geografia y Meteoro- 
logico Nacional, Mexico D.F. 

ANON. 1981. Carta de uso del suelo y vegetacion 1:1,000,000. Mexico. Estados Unidos 
Mexicanos, Mexico, Guadalajara and Mexico. Direccion General de Geografia del 
Territorio Nacional. San Antonio de Abad 124, Mexico 8, D.F. 

ARBINGAST, S. A., C. P. BLAIR, J. R. BUCHANAN, C. C. GILL, R. K. HOLZ, C. A. MARIN 
R., R. H. RYAN, M. E. BONINE & J. P. WEILER. 1975. Atlas of Mexico. Bureau of 
Business Research, University of Texas, Austin. 165 pp. 

BARKER, J. F. & W. S. HERMAN. 1976. Effect of photoperiod and temperature on 
reproduction of the monarch butterfly, Danaus plexippus. J. Insect Physiol. 22:1565— 
1568. 

BATSCHELET, E. 1972. Recent statistical methods for orientation data. Pp. 61-69 in S. 
R. Galler, K. Schmidt-Koenig, G. T. Jacobs & R. E. Belleville (eds.), Animal orien- 
tation and navigation. N.A.S.A., Washington, D.C. 606 pp. 

BREWER, J. 1983. The reign of the monarchs. Geo 5:92-97. 

BROWER, L. P. 1985. New perspectives on the migration biology of the monarch 
butterfly, Danaus plexippus L. Pp. 748-786 in M. A. Rankin (ed.), Migration: Mech- 
anisms and adaptive significance. University of Texas Marine Science Institute. Con- 
tributions to Marine Science, Supplement 27. 867 pp. 

In Press. Biological rationale for the conservation of the overwintering sites of 
the monarch butterfly, Danaus plexippus L., in the states of Michoacan and Mexico. 
In: Proc. First International Wildlife Symposium, Mexico City. 

BROWER, L. P. & W. H. CALVERT. 1985. Foraging dynamics of bird predators on 
overwintering monarch butterflies in Mexico. Evolution 39:852-868. 

BROWER, L. P., W. H. CALVERT, L. E. HEDRICK & J. CHRISTIAN. 1977. Biological 
observations on an overwintering colony of monarch butterflies (Danaus plexippus 
L.) in Mexico. J. Lepid. Soc. 31:282-242. 

BROWER, L. P., B. E. HORNER, M. M. Marry, C. M. MoFrFIT & B. VILLA R._ 1985. 
Mice (Peromyscus maniculatus labecula, P. spicelegus, and Microtus mexicanus 





186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


salous) as predators of overwintering monarch butterflies (Danaus plexippus) in 
Mexico. Biotropica 17:89-99. 

CALVERT, W. H. & L. P. BROWER. 1981. The importance of ground escape behavior 
and forest cover for the survival of overwintering monarch butterflies, (Danaus 
plexippus, Danaidae). J. Lepid. Soc. 35:216—225. 

CALVERT, W. H. & J. COHEN. 1983. The adaptive significance of crawling up onto 
foliage for the survival of grounded overwintering monarch butterflies (Danaus 
plexippus) in Mexico. Ecol. Entomol. 8:471-—474. 

CALVERT, W. H., L. E. HEDRICK & L. P. BROWER. 1979. Mortality of the monarch 
butterfly (Danaus plexippus L.): Avian predation at five overwintering sites in Mex- 
ico. Science 204:847-851. 

CALVERT, W. H., M. B. HyatT & N. P. MENDOzA V. 1987. The effects of understory 
vegetation on the survival of overwintering Monarch butterflies (Danaus plexippus 
L.) in Mexico. Acta Zool. Mexicana (in press). 

CALVERT, W. H., W. ZUCHOWSKI & L. P. BROWER. 1982. The impact of forest thinning 
on microclimate in monarch butterfly (Danaus plexippus L.) overwintering areas of 
Mexico. Bol. Soc. Bot. Mexico 42:11-18. 

1983. The effect of rain, snow and freezing temperatures on overwintering 
monarch butterflies in Mexico. Biotropica 15:42—47. 

CASTELLANOS, P. 1983. La mariposa monarca. Documentary produced by Channel 13 
Television, Mexico, D.F. 

CHAPLIN, S. B. & P. H. WELLS. 1982. Energy reserves and metabolic expenditure of 
monarch butterflies overwintering in Southern California. Ecol. Entomol. 7:249-256. 

DE LA Maza, R. 1976. La mariposa y sus estilizaciones en las culturas Teotihuacana 
(200 a 750 D.C.) y Azteca (1325 a 1521 D.C.). Rev. Soc. Mex. Lepid. 2(1):39-48. 

DE LA MAZa, R. G., J. DE LA MAZA & A. DiAZ FRANCES. 1977. Movimientos migratorios 
de “Monarcas” en el Estado de Oaxaca, Mexico. Bol. Inform. Soc. Mex. Lepid. A.C. 
3(5):12-18. 

GEIGER, R. 1950. The climate near the ground. Harvard University Press, Cambridge, 
Massachusetts. 482 pp. 

GOLDMAN, E. A. 1951. Biological investigations in Mexico. Smiths. Misc. Coll. 115:1- 
476. 

GOLDMAN, E. A. & R. T. Moore. 1946. The biotic provinces of Mexico. J. Mammal. 
26:347-360. 

KAMMER, A. E. 1970. Thoracic temperature, shivering, and flight in the Monarch 
Butterfly, Danaus plexippus (L.). Z. Vergl. Physiol. 68:334-344. 

KIEMELE Muro, M. 1975. Vocabularia Mazahua-Espanol y Espanol Mazahua. Edicion 
preparada por M. Colin. Biblioteca Enciclopedica del Estado de Mexico, Mexico D.F. 
447 pp. 

MALCOLM, S. B., B. J. COCKRELL & L. P. BROWER. 1987. Monarch butterfly voltinism: 
effects of temperature constraints at different latitudes. Oikos (in press). 

Masters, A. R. 1985. The influence of temperature on the overwintering biology of 
the monarch butterfly (Danaus plexippus). MS Thesis, Univ. of Florida, Gainesville. 
88 pp. 

MookreE, R. T. 1945. The transverse volcanic biotic province of central Mexico and its 
relationship to adjacent provinces. Trans. San Diego Soc. Nat. Hist. 10:217-236. 
MOsINO ALEMAN, P. A. & E. GarciA. 1974. The climate of Mexico. Pp. 345-404 in 

R. A. Brysen & F. K. Hare (eds.), World survey of climatology. Elsevier, Amsterdam. 

RZEDOWSKI, J. 1978. Vegetacion de Mexico. Editorial Limusa, Mexico, D.F. 432 pp. 

SANCHEZ, S., O. 1979. La flora del valle de Mexico. Editorial Herrero, S.A. 519 pp. 

SANTAMARIA, F. J. 1974. Diccionario de Mejicanismos. 2nd ed. Editoria Porrua S. A., 
Mexico. 1207 pp. 

SCHMIDT-KOENIG, K. 1979. Directions of migrating monarch butterflies (Danaus plex- 
ippus; Danaidae; Lepidoptera) in some parts of the eastern United States. Behav. 
Proc. 4:73-78. 

TUSKES, P. M. & L. P. BROWER. 1978. Overwintering ecology of the monarch butterfly, 
Danaus plexippus L., in California. Ecol. Entomol. 3:141-153. 





VOLUME 40, NUMBER 3 187 


UrQuuHART, F. A. 1960. The monarch butterfly. University of Toronto Press, Toronto, 
Canada. 361 pp. 

1976. Found at last: The monarch’s winter home. Nat. Geogr. 150:160-173. 

URQUHART, F. A. & N. R. URQUHART. 1978. Autumnal migration routes of the eastern 
population of the monarch butterfly (Danaus p. plexippus L.; Danaidae: Lepidop- 
tera) in North America to the overwintering site in the Neovolcanic Plateau of 
Mexico. Can. J. Zool. 56:1754-1764. 

WEsT, R. C. 1964. Handbook of the Middle American Indians, Vol. 1. University of 
Texas Press, Austin. 570 pp. 

YAMAGUCHI, H. 1980. TV Production on the Monarch Butterfly, Nippon TV Corpo- 
ration, Tokyo, Japan, 28 February 1980. 

ZEPEDA, T. 1941. La Republica Mexicana Geografia y Atlas. Editorial Progreso Mexico, 
D.F. 152 pp. 





Journal of the Lepidopterists’ Society 
40(3), 1986, 188-190 


A NEW EUCHLOE (PIERIDAE) FROM 
NORTHWESTERN MEXICO 


PAUL A. OPLER 


1025 Pennock Place, 
Fort Collins, Colorado 80524 


ABSTRACT. A new species of Euchloe related to Euchloe ausonides was discovered 
at a mountainous desert scrub locality in Sonora, Mexico. This butterfly is described from 
four males and compared with Euchloe ausonides, its close relative; no females are 
known. Its habitat and evolutionary significance are treated. 


Before the discovery reported here, four species of Euchloe Hubner 
were known from the Americas—the most recently described being E. 
hyantis and E. olympia, both described by William Henry Edwards 
in 1871. More recent study has elucidated the systematic position, dis- 
tribution, early stages, and biological features of these four species 
(Opler, 1966a, b, c, d, 1970, 1971, 1974, Opler & Clench 1983). 

The discovery of a new Euchloe species by Richard Holland of 
Albuquerque, N.M. in northwestern Mexico was quite unexpected. The 
species is a member of Ausonides Species Group. 


Euchloe guaymasensis Opler, new species 


Male. Forewing length 19.0 mm (range 17.4-19.4 mm, N = 5). Head. Labial palpi 
covered with white appressed scales, long stiff white and black hairlike setae directed 
laterally and ventrally; frons with long white and black setae; vertex with mixture of 
yellow and a few black appressed scales; antennae 0.07 cm long, white-tipped, otherwise 
covered with a mixture of white and black scales, predominantly the former, inner 
surface of shaft and nudum naked. 

Thorax. Clothed dorsally with quadrate black appressed scales and fine long white 
hairlike setae; laterally and ventrally clothed with yellow appressed scales together with 
long white hairlike scales; legs covered mainly with white and a few black scattered 
scales. Forewing. Apex rounded with stem R4+5 much shorter than R5; dorsal surface 
ground pale lemon yellow, with white along costal margin from base to apex; white 
diffusely along inner margin; very fine white hairlike scales especially at base and along 
inner margin; black patch at end of discal cell concave outwardly, 9 scale rows in width 
with 12-14 included scattered white scales; black patch at apex dense with few included 
yellow scales; oblong patch (rounded in several paratypes) of pale yellow scales on costal 
margin between R1 and R3; fringe at apex black except for three areas with long yellow 
hairlike setae; few scattered appressed black scales at base; ventral surface pale yellow; 
small greenish patches (composed of appressed yellow and black scales) where R1 meets 
costal margin and where M2 and M3 meet outer margin; patch at end of discal cell of 
black scales with included gray white scales. Hindwing. Dorsal surface ground pale 
lemon yellow, some white along inner margin, a few appressed black scales at base; long 
fine pale hairlike scales especially on base and along inner margin; a few (5-15) scattered 
black appressed scales at endings of veins (except SC and Anals); fringe of long fine 
yellow hairlike scales except for a few long black scales at vein endings as above. Ventral 
surface ground white; marbling pattern similar to that of E. ausonides but with fewer 
connections between patches of yellow and black scales; very long (0.6 cm) “green” 
patch within and parallel to interior margin of discal cell. Abdomen. Covered dorsally 
and laterally with black appressed scales, a few scattered yellow or white scales inter- 


VOLUME 40, NUMBER 3 189 





Fic. 1. Holotype male Euchloe guaymasensis, dorsal view. 
Fic. 2. Paratype male E. guaymasensis, ventral view. 


Fic. 3. Euchloe guaymasensis male genitalia, lateral view. 


mixed; ventrally with mixture of quadrate white and yellow appressed scales, covered 
with medium length white hairlike scales. Genitalia. As in Euchloe ausonides except 
saccus more regular in outline; cucullus areas of valvae only slightly produced after distal 
teeth; juxta only weakly chitonous, only dorsal arms visible. 

Types. Holotype 6, Mexico, Estado de Sonora, Las Avispas microwave relay, 2000’ 
(655 m), 40 mi (64 km) N Guaymas, 24 March 19838, leg. Richard Holland; genitalia 
preparation PAO 354 (P. A. Opler). Paratypes. Same locality, 4 66; 12 March 1974, 23 
March 1983 (2), 5 March 1984; all leg. Richard Holland (the 1974 paratype with G. S. 
Forbes). Holotype deposited in National Museum of Natural History, Smithsonian Insti- 
tution, Washington, D.C. One paratype each in collections of American Museum of 
Natural History, New York; Allyn Museum of Entomology, University of Florida; Los 
Angeles County Museum; Instituto de Biologica, Universidad Nacional Autonoma de 
Mexico. 

The name is masculine and refers to Guaymas, the nearest city to the type locality. 


COMPARATIVE ANALYSIS 


Among the four previously described Euchloe, E. guaymasensis is 
most closely allied to E. ausonides (Lucas); the two species share the 
following characters: (1) antennae with black and white scales, (2) 
white scattered scales within forewing dorsal discal cell black patch, 
(3) lack of opalescent white scales on ventral hindwing, (4) marbling 
pattern on ventral hindwing neither heavy and fractured nor strongly 
reduced, (5) cucullus extending slightly beyond distal tooth, (6) juxta 
V-shaped, and (7) sterigma evenly curved in lateral view. Opler (1971) 
illustrated these character states, some of which—but not all—are also 
shared by either E. creusa (Doubleday) or E. olympia (Edwards) or 
both. Compared to Euchloe ausonides, E. guaymasensis differs in the 
following features: (1) antennae with a greater proportion of white 
scaling, (2) forewing more rounded at apex, (8) black patch at apex 
darker, without intermixed white scales, (4) forewing costal margin 
without black scaling, (5) ground color of both dorsal wing surfaces 
pale lemon yellow (usually cream white in E. ausonides), (6) forewing 
stem R4+5 much shorter than R5, (7) black scaling areas at bases of 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


both wings much more restricted, (8) green marbling patches on ven- 
tral hindwing less interconnected, and (9) male genitalia with juxta less 
chitonous. 

Habitat at type locality: photographs taken by Richard Holland at 
the type locality show rocky, extremely rugged, mountainous terrain 
with a mixture of subshrubs, succulents, and woody microphyllous 
deciduous vegetation. Specific plants include ocotillo (Fourquieria), 
Acacia, and possibly Encelia. The type locality is clearly within the 
Lower Sonoran life zone. 

Surprisingly, the closest nearctic relative of Euchloe guaymasensis 
is E. ausonides, a species of much more temperate climes, not E. 
hyantis, a butterfly of more austral habitats, including northern Sonora, 
Mexico. The closest localities to the E. guaymasensis type locality where 
E. ausonides occurs are the Kaibab Plateau, Coconino Co., Arizona, 
975 km to the north, and the Sangre de Christo Mountains, San Miguel 
Co., New Mexico, 965 km to the northeast. 

Likely it will be found elsewhere in the coastal ranges of Sonora 
when someone penetrates this physically intimidating habitat. 

Euchloe guaymasensis differs in sufficient characters from E. au- 
sonides to indicate that it has been long isolated, probably since at least 
the middle Pleistocene. The nature of the larval and pupal characters, 
when discovered, should help confirm this hypothesis. 


ACKNOWLEDGMENTS 


I thank Richard Holland for allowing me to describe this new species. The U.S. Fish 
and Wildlife Service, Denver Wildlife Research Center, Branch of Ecology provided 
laboratory facilities. John Buffington, Colorado State University, exhibited skill and pa- 
tience in providing the photographs. Robert K. Robbins and an anonymous reviewer 
provided suggestions that improved the manuscript. 


LITERATURE CITED 


OPLER, P. A. 1966a. Studies on the Nearctic Euchloe. Part 1. Introduction. Jour. Res. 
Lepid. 5:39—40. 

1966b. Studies on the Nearctic Euchloe. Part 2. Chronological review of the 

literature and bibliography. Jour. Res. Lepid. 5:41-50. 

1966c. Studies on the Nearctic Euchloe. Part 3. Complete synonymical treat- 

ment. Jour. Res. Lepid. 5:185-190. 

1966d. Studies on the Nearctic Euchloe. Part 4. Type data and type locality 

restrictions. Jour. Res. Lepid. 5:190-195. 

1970. Studies on the Nearctic Euchloe. Part 5. Distribution. Jour. Res. Lepid. 

7:65-86. 

1971. Studies on the Nearctic Euchloe. Part 6. Systematics of adults. Jour. Res. 

Lepid. 8:153-168. 

1974. Studies on the Nearctic Euchloe. Part 7. Comparative life histories, hosts 
and the morphology of immature stages. Jour. Res. Lepid. 13:1-20. 

OpLER, P. A. & H. K. CLENCH. 1983. Studies on the Nearctic Euchloe. 8. Euchloe 
olympia. Ann. Carnegie Mus. 52:41-54. 




















Journal of the Lepidopterists’ Society 
40(3), 1986, 191-205 


PHYSICAL CONSTRAINTS OF DEFENSE AND RESPONSE TO 
INVERTEBRATE PREDATORS BY PIPEVINE CATERPILLARS 
(BATTUS PHILENOR: PAPILIONIDAE) 


NANCY E. STAMP 


Department of Biological Sciences, State University of New York, 
Binghamton, New York 13901 


ABSTRACT. The responses of pipevine swallowtail caterpillars (Battus philenor: 
Papilionidae) to simulated attacks of invertebrate enemies and to actual attack by coc- 
cinellid larvae (Hippodamia convergens: Coccinellidae) were examined. The caterpillars 
were more reactive to the simulated attack of a biting predator than to the simulated 
touch by an insect enemy. Active fifth instars reached around to the posterior or walked 
away in response to stimuli, whereas prepupal fifth instars were more likely to extrude 
the osmeterium and never moved away from the stimuli. Caterpillars that were larger 
than the coccinellid predators were attacked but seldom eaten. In contrast, larvae that 
were the same size or smaller than the coccinellids were killed more frequently. When 
the caterpillars were attacked posteriorly, they defended a limited area by reaching 
around while the prolegs remained attached. The area defended depends on cuticular 
stretch, number of attached prolegs, current physiological state, and type and degree of 
stimulation. 


A common view of insects as prey is that behaviorally they are rather 
defenseless. Dixon (1973) stated, “The general impression conveyed by 
the literature is that aphids and related small insects are helpless, sed- 
entary and thin-skinned creatures that invite the attention of any pred- 
ator that comes along.’’ Generally, that same view is held for caterpil- 
lars and other immature insects. Yet caterpillars can and will defend 
themselves under certain circumstances, such as when attacked by 
insect predators and parasitoids. However, as I show here, pipevine 
swallowtail caterpillars (Battus philenor (L.): Papilionidae) have im- 
portant constraints that limit the effectiveness of defensive behavior. 

Most six-legged insects can turn up to 360° in their own defense. For 
instance, aphids can move forward or backward rapidly, and they can 
kick their adversaries (Banks 1957). Aphids can escape just before con- 
tact or immediately afterward by simply moving away quickly; many 
invertebrate predators must be within a centimeter of, or bump into, 
prey before detecting them (Russel 1972). Furthermore, before or after 
contact by predators, aphids can swivel on their stylets up to 180° and 
thus continue feeding while avoiding their enemies (Russel 1972, Brown 
1974). In contrast, caterpillars have a cylindrical body with short tho- 
racic legs anteriorly and short prolegs posteriorly. This means that they 
differ greatly in their maneuverability from six-legged insects, espe- 
cially aphids which are pear-shaped with relatively long legs. As a 
consequence of the mobility of six-legged insects, most invertebrate 
predators are more mobile than caterpillars and can outmaneuver them. 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Thus, caterpillars may benefit by escape, often dropping off their 
host plant (Myers & Campbell 1976). However, leaving the host plant 
can be risky. Caterpillars (and other soft-bodied insects) may starve 
before they locate a host plant, or die from desiccation or ground 
predation (Dethier 1959, Roitberg & Myers 1978, Rausher 1979). Lar- 
vae may drop on a thread of silk, which they can later climb, returning 
safely and quickly to their host plant (Dempster 1971). But predatory 
ants may climb down silk threads to capture larvae (Allen et al. 1970). 

Some caterpillars feed in hiding, a strategy that appears to be es- 
pecially effective for early instars (Allen et al. 1970, Lopez et al. 1976). 
Web-making caterpillars may disappear into their webs when dis- 
turbed (Fitzgerald 1980). But webs do not deter some wasps and pen- 
tatomids from pursuing caterpillars within (Morris 1972, Schaefer 1977). 
Carabid beetles may tear open webs (Langston 1957). Chrysopid larvae 
with their long, sicklelike mandibles, and pentatomid and reduviid 
bugs with their long beaks, can attack prey through cloth and webbing 
(Fleschner 1950, Bornemissza 1966, Allen et al. 1970, Berisford & Tsao 
1975). Furthermore, some predatory pentatomids and spiders live in 
webs of caterpillars (Morris 1972, E. W. Evans, pers. comm.). 

Therefore, caterpillars may benefit by vigorous defense when escape 
is less effective or more risky, such as when an insect predator initiates 
contact but cannot overwhelm the caterpillar. Typically, a defensive 
caterpillar attaches firmly to the substrate with the prolegs, lifts the 
thoracic legs and swings the anterior of the body toward the attacker, 
especially when approached from the side or rear by a predator. Cat- 
erpillars may use their bodies to hit and their mandibles to grasp an 
attacker (Morris 1963, McFadden 1968, Iwao & Wellington 1970, Frank 
1971, Heinrich 1979, Suzuki et al. 1980, Stamp 1982). Unlike verte- 
brates and adult insects, caterpillars do not use their legs defensively. 
Instead, they may regurgitate or wipe offensive glands on attackers 
(Eisner & Meinwald 1965, Feltwell 1982). 

The questions posed in this study were: 1) when does a caterpillar 
opt to escape or for defense? 2) how does it defend itself? and 3) how 
effective is it in defending itself, or when are insect enemies successful 
in countering a caterpillar’s defense? 


METHODS 


Pipevine swallowtail caterpillars were used because of their variety 
of defensive responses: thrashing with the front half of the body, grasp- 
ing with the mandibles, regurgitating, and extruding the osmeterium 
and wiping it on attackers. On 16 May 1983, eggs were collected at 
Rancho Cordova, California, along with Dutchman’s pipevine (Aris- 
tolochia californica Torr.), the host plant. Caterpillars were reared in 


VOLUME 40, NUMBER 3 193 


the laboratory at room temperature in plastic boxes with the host plant 
stems in aquapics. 


Response to Simulated Attack 


To examine the responses by swallowtail caterpillars, I used two 
stimuli. On first, second, third, and fifth instars, a two-haired brush 
simulated the touch of an insect predator (palpitating antennae or beak 
of predatory hemipteran) or parasitoid (palpitating antennae or prob- 
ing ovipositor). The stimulus was applied three consecutive times to 
the posterior of each caterpillar at 15 sec intervals for up to 6 trials. 
For the fifth instar, a pinch with forceps on a fleshy tubercle at the 
rear end at 15 sec intervals simulated the bite of an insect predator, 
such as an ant. Responses were recorded with a video camera. Thrash- 
ing with the front half of the body, biting with the mandibles, extrud- 
ing the osmeterium, and regurgitating were classified as stationary 
defense. Walking away and wriggling (rolling around with no legs 
attached) were classified as escape behaviors. For Chi-square analyses, 
the Yates correction for continuity was used when v = 1, and is denoted 
by x2 (Zar 1974). The power of tests (probability of not committing a 
type II error, 1 — 8) was calculated as described by Cohen (1977). 

To compare active and inactive larvae, the stimuli were applied in 
the same manner as above to two sets of final instars: feeding and 
prepupal (no longer feeding and residing on a silk mat). 


Tests with Invertebrate Predator 


To determine how effective swallowtail caterpillars were in defend- 
ing themselves, they were tested with coccinellid larvae (Hippodamia 
convergens Guerin-Meneville). Coccinellid larvae are voracious pred- 
ators of eggs and small insects (Banks 1957, Brown 1974). I observed 
a third instar H. convergens feeding on a second instar B. philenor in 
a riparian area in Rancho Cordova; it is thus reasonable to assume that 
the coccinellids are natural predators of these swallowtail caterpillars. 
Fourteen third instar coccinellids were collected on 23 May 1983 at 
Rancho Cordova, California, in a large field that had no Battus phi- 
lenor caterpillars or host plants. Thus, the predators would not have 
had any prior contact with pipevine swallowtail caterpillars. 

Each coccinellid was kept in a Petri dish with a source of water but 
no food for 24 h. Using a paintbrush, I placed each coccinellid in a 
Petri dish with a swallowtail caterpillar on a leaf of the host plant. 
Caterpillars were used only if they were actively feeding. Fluon (poly- 
tetrafluoroethylene from Imperial Ltd., England) was painted on the 
sides and bottom of the dish to prevent the coccinellids from searching 
there. The interactions of each coccinellid and caterpillar were moni- 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


tored for 10 min. Only those trials where the coccinellids exhibited 
foraging behavior were used for analysis. Some coccinellids were used 
more than once but only at 24-h intervals. To determine the body-size 
ratio (BSR), body length of caterpillars was measured in mm when 
they were eating and when the dorsal midline (anterior to posterior) 
was straight. Body length of the coccinellids was measured when they 
were between foraging bouts and thus motionless. 


Modelling Defensive Space 


The area defended by swallowtail caterpillars was determined by 
using a video camera to record the path of the head in response to 
stimuli (two-haired brush or forceps) applied to the posterior. Mea- 
surements were made from tracings of body length at the start, and 
how close the head came to the posterior during defense. Using a map 
meter, measurements were also made of body length (down the mid- 
line) when the caterpillar reached around to defend itself. Body exten- 
sion was estimated by maximal body length during defense divided by 
body length at the start. Only the recordings where the head moved 
laterally rather than over the back were analyzed. While the camera 
recorded body movement, I recorded the number of prolegs detached 
immediately after the stimulus. 


RESULTS 
Response to Simulated Attack 


In response to the two-haired brush (simulating the touch of an 
invertebrate predator or parasitoid), all of the tested instars exhibited 
escape and defensive behaviors (Fig. 1). With repeated stimulation, 
the larvae were more likely to exhibit escape behaviors, except for the 
fifth instar where the escape response declined. Defensive responses 
decreased with the sequence of stimuli. The fifth instar appeared to 
become more tolerant of the stimuli in that both defense and escape 
responses declined with the sequence of stimuli (Fig. 1). 

Comparing responses of the fifth instar to the two stimuli showed 
that they were more reactive, by exhibiting responses, to pinching than 
to touching by the brush (x2, P < 0.025). But the larvae responded 
with escape and defensive behaviors in similar proportions (escape, 
defense or no response; x2 test, v = 2, P > 0.10, 1 — B = 0.58 for a = 
0.05, n = 58). They were more likely to reach backward in response 
to the forceps than to the brush (x2, P < 0.025). The caterpillars also 
walked farther away in response to pinching than to touching by the 
brush (moving more than 2 cm within 15 sec or not, x2, P < 0.05). 

Comparisons were made of behavior of active (still feeding) and 


VOLUME 40, NUMBER 3 195 


Percent of 


larvae 


100 
A. Defense 


50 








12253-45516 102-34, 5:6 172/34 5°56 





100 ie 


B. Escape 


90- 





123456 123456 123456 123 


ie I2 I3 ES 
Series of responses 


Fic. 1. Reaction by swallowtail caterpillars to a two-haired brush touching the pos- 
terior at intervals of 15 sec (indicated by the series of numbers on the x-axis). Sample 
sizes were n = 9 for the first and second instars (I1, I2), n = 11 for the third instar (I3) 
and n = 30 for the fifth instar (I5). Responses were classified as defense or escape. Some 
larvae exhibited both types of response at a single trial. A. Defensive responses. B. Escape 
behaviors. 


inactive (prepupal and no longer feeding) fifth instars. In contrast to 
active larvae, prepupal caterpillars exhibited no escape behavior. They 
responded defensively to the brush in the same proportion as caterpil- 
lars still feeding (x2, P > 0.50, 1 — 6 = 0.12 for a = 0.05, Fig. 2). 
However, prepupal larvae were more likely to extrude the osmeterium 
than feeding caterpillars (x2, P < 0.025). Although both the active 
(feeding) and inactive (prepupal) larvae reached around or thrashed, 
the active larvae were able to reach to their posterior or to walk away, 
whereas the prepupal caterpillars were much less mobile and never 
walked away. 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PeSreenr Or 
larvae 


100 
ie E<iinwele 


Reach 
a Both 





\ 2eS Were ome Li 2 M2 Seema 


LL] PP | PP 
Brush Pinch 


Fic. 2. Responses by active (LI—still feeding) and inactive (PP—prepupal and no 
longer feeding) final instar swallowtails to either a brush or forceps. Numbers on the 
x-axis represent the sequence of stimuli applied at intervals of 15 sec. Sample sizes were 
30 and 28 for the active larvae stimulated by brush and forceps, respectively, and 35 for 
the prepupal larvae. Extrude—osmeterium extruded, with an attempt by the caterpillar 
to wipe them on the brush or forceps. Reach—caterpillar reached with head backwards 
toward stimulus. 


Tests with Coccinellid Predators 


To examine effectiveness of defense, caterpillars were monitored in 
response to invertebrate predators (coccinellid larvae). When the cat- 
erpillars were the same size or smaller than the predators (n = 16), 
43.8% of the caterpillars were eaten. In contrast, when the caterpillars 
were larger than the predators (n = 16), only 6.8% were eaten. The 
frequencies of eaten and uneaten caterpillars were significantly differ- 
ent (x2, P < 0.05). Examination of the behavior of these caterpillars 
showed that large caterpillars (body-size ratio of prey to predator great- 
er than 1) were more likely to thrash in response to the coccinellids, 
and small caterpillars (relative to the predators) were more likely to 


VOLUME 40, NUMBER 3 197 


PERCENT OF 
LARVAE 
100 


LI esr < 10 
BSR > 1.0 





Move Wriggle Thrash Bite Extrude 
away 


RESPONSE 
Fic. 3. Reaction by swallowtail caterpillars to third instar coccinellid predators. Re- 
sponses are divided into those where body-size ratio (BSR) of caterpillar to predator was 
greater than 1, or less than or equal to 1. 


wriggle (no legs attached) (x? tests, P < 0.001, Fig. 3). In this experi- 
ment, body size ratio had no effect on escape, biting, or on extruding 
the osmeterium (Fig. 8). 

Third instar coccinellids had little difficulty subduing first instar 
swallowtails, which were smaller than the predators. For example, after 
a coccinellid touched one of six first instars feeding together, the touched 
caterpillar extruded its osmeterium. The predator backed off, then 
approached and bit the larva behind its head. The other larvae had 
been feeding and moving around. After the first caterpillar was at- 
tacked, the rest remained motionless for 11 min. Only after the coc- 
cinellid backed into one of them did the rest leave the area. Although 
it took up to 15 min for a coccinellid to consume a first instar, these 
predators ate as many as three caterpillars consecutively. All of the 
coccinellids that ate pipevine swallowtail caterpillars survived and pu- 
pated, which suggests that these caterpillars are appropriate prey for 
the coccinellids. 


198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





° 
SS o Slovo) fo oe 


v 
V4 


< i > 

Fic. 4. Example of typical movement of a final instar swallowtail larva in response 
to pinching stimuli (applied at right), based on tracing body movement from a video 
record. 1—position of head just before first stimulus on posterior. Open triangles show 
path of head after first stimulus. 2—position just before second stimulus, applied 15 sec 
after the first. Dark triangles indicate path of head after second stimulus. 


When predator and caterpillar were of similar size, caterpillars were 
usually successful in defending themselves. For instance, after contact 
by a coccinellid, one caterpillar thrashed vigorously. The predator 
grabbed it. The caterpillar responded by biting the legs of the cocci- 
nellid but did not use its osmeterium. The coccinellid released the 
caterpillar, moved away, and began grooming. 


Modelling Defensive Space 


To determine how capable larvae were at defending their posteriors, 
the path of the head was traced using video recordings of larval re- 
sponse. As shown in Figs. 4 and 5, the caterpillars defended a limited, 
circular area around themselves. This was a consequence of their head- 
to-rear defense, where they did not turn around and place their pos- 
terior away from the stimulus as most animals do. Both stimuli elicited 
the head-to-rear response. Although some caterpillars may feed while 
attached to a branch, and consequently are restricted in their ability 
to turn around, many caterpillars are free to move around on the leaves 
on which they reside. Those in this study were unrestricted also. Thus, 


VOLUME 40, NUMBER 3 199 


BEAD: TO 
MIDLINE IN CM 
O25) 

Gon ee! EAD AT MIDUINE | 2722 2 gee!) 
| v 
gia 

OO : re 
| 

-1.0 os a rok 

vi A J 
Vv Vv a 
Vv ee, 
ey i) A < 
Vv rely a) 
A a 4 < 
uJ 
42K) a 
A Vv | 
y | 
mel 





MOO ace 7:20 sale THOM Pay aks) OO Ret OS LO 
AND SM OMiRNESATR. niIN Ts CM 


Fic. 5. Response by fifth instar swallowtails to either the two-haired brush or forceps 
applied to the posterior (arrow). Caterpillar’s head always began at the top left and 
moved to the side (down and to the right). The farthest point of the head back toward 
the posterior is shown. Open triangles indicate response to the brush by 11 caterpillars, 
dark triangles the response to the forceps by 20 caterpillars. 


these caterpillars exhibited a head-to-rear defense even when they could 
have turned around completely to face attack. 

Of 31 fifth instars, 48% responded by reaching to or beyond the 
posterior. That so many did so was due to two factors: body extension, 
and detaching some prolegs. In response to the brush, body extension 
(beyond the original length) was 30% (+4.0 SE, n = 11). Extension was 
33% (+4.5 SE, n = 20) in response to the forceps, with no significant 
difference between the two stimuli (arcsin transformation, two-sample 
t-test, P > 0.50, df = 1,29, 1 — B = 0.46 for a = 0.05). These caterpillars 
have five pairs of prolegs but may detach up to four of the anterior 
pairs in defending themselves. The mean number of attached pairs 
after brush stimulation was 4 (+0.3 SE, n = 11), and after pinching, 4 
(+0.1 SE, n = 20, square-root transformation, two-sample t-test, P > 
0.20, df = 1,29, 1 — 6 = 0.46 for a = 0.05). 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


DISCUSSION 
Response to Attacks 


Defensive and escape behaviors varied with instar and physiological 
state (active versus prepupal larvae). For example, the prepupal larvae 
were more likely to extrude the osmeterium than active caterpillars 
(those still feeding). The active larvae were mobile and thus could 
escape, whereas the prepupal larvae were not mobile, were slow to 
respond, and thus had no escape options and few defensive ones. Cor- 
respondingly, invertebrate predators and parasitoids may be more suc- 
cessful when attacking relatively defenseless (inactive or small) insects 
than when attacking active and larger ones (Dixon 1959, Evans 1976, 
Cate et al. 1977, Tilman 1978). For example, Rabb and Lawson (1957) 
stated that “an appreciable number” of tobacco hornworms captured 
by Polistes wasps were molting, and often those wasps approached but 
left alone fifth stage (large) larvae. Iwao and Wellington (1970) found 
that tent caterpillars differed in their behavior, with inactive types 
being less defensive and more often parasitized. Similarly, in this study 
coccinellid predators were more successful in their attacks when B. 
philenor caterpillars were smaller than they were. 

The caterpillars defended themselves by biting the coccinellids, par- 
ticularly on their legs, by extruding the osmeterium and wiping it on 
the coccinellids, and occasionally by thrashing and regurgitating. The 
caterpillars were more reactive to pinching than to touch by the brush. 
Probably pinching provided a clear signal of attack, whereas the brush 
stimulus may be received as a more general signal and not clearly 
different from the touch of a leaf moved by the wind. Relatively large 
caterpillars thrashed in response to the coccinellids; most of the time 
the coccinellids responded by backing off and eventually leaving the 
caterpillars alone. In contrast, the small larvae often wriggled when 
touched by the coccinellids. By wriggling, the caterpillars moved er- 
ratically, which may have made it difficult for the predators to respond 
effectively to the prey (Humphries & Driver 1967); some caterpillars 
rolled off the leaf and escaped from predators. 


Physical Constraints of Defense 


The premise here is that by moving only the front half of the body, 
the typical caterpillar defends a limited space around itself (to either 
side and over its back, Fig. 6). Variables that affect the maximal de- 
fendable area are: L, the length of the moving (defending) portion of 
the body; D, the diameter of the body; and M, the length of the moving 
portion of the body after maximal cuticular stretch. To estimate M, 
the cuticle was extended by 25% (a reasonable estimate based on Hep- 


VOLUME 40, NUMBER 3 201 





Fic. 6. A. Defendable area of a typical caterpillar when arching the head over its 
back. B. Shaded portion shows maximal defendable area when there is no cuticular 
stretch, dashed line when there is cuticular stretch. C. With only two pairs of prolegs 
attached, a caterpillar can defend a much larger area than depicted in A. 


burn & Levy 1975, Wolfgang & Riddiford 1981, Fig. 6B). Cuticular 
stretch lengthens the outer bending side and thus accommodates the 
body volume. The effect of cuticular stretch is to increase the reach 
and, consequently, maximal defendable area of the caterpillar. The 
illustrations (Figs. 4, 5 & 6) indicate where caterpillars may be most 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


vulnerable to attack by insect enemies that cannot simply overwhelm 
them. That area is some portion of the rear end. With some prolegs 
detached, a caterpillar can defend a larger area and reach beyond the 
posterior than when five sets of prolegs are attached (Fig. 6A & C). 
Thus, the maximal defendable area of a particular caterpillar should 
affect: 1) type of defensive behavior employed by the caterpillar; 2) 
attack behavior of enemies, especially specialist parasitoids; 3) the se- 
quence of attack-defense behaviors exhibited during enemy-caterpillar 
interactions; and 4) outcome of the event. 

Using measurements of body diameter and the moving (defending) 
portion of a caterpillar relative to total body length, the maximal de- 
fendable space can be estimated (Fig. 6). When the ratio of the moving 
portion relative to total body length is high, or when diameter of the 
body is small, the caterpillar can reach its posterior easily (Fig. 6C). A 
slender geometrid caterpillar that has two pairs of prolegs uses this 
increased maneuverability to its advantage by preying on small insects 
that touch its posterior (Montgomery 1982). Cuticular elongation may 
also affect maximal defendable space because it allows cuticular ad- 
justment (stretch of the outer bending surface) to accommodate a body 
volume that is more or less compressible, depending on how close the 
caterpillar is to molting to the next instar (Fig. 6B). 

Most invertebrate predators rely on tactile and chemical cues more 
than visual ones to detect prey (chrysopid larvae, coccinellid larvae 
and adults, predatory wasps, pentatomid bugs; Banks 1957, Klingauf 
1967, Fleschner 1950, Storch 1976, Steiner 1974, Hicks 1931, Evans 
1982). Usually, such predators encounter their prey physically before 
they attack (Banks 1957, Dixon 1959, Swynnerton 1915, Allen et al. 
1970, Myers & Campbell 1976). Consequently, predators may benefit 
by attacking whatever part of the prey they encounter and thus pre- 
vent the victim’s escape (Brown 1974, Evans 1982). 

But invertebrate predators and parasitoids often attack prey and 
hosts cautiously. For instance, predatory pentatomids (Perillus circum- 
cinctus) tested most potential prey (chrysomelid larvae) by extending 
the beak and then retreating when the prey defended themselves (E. 
W. Evans, pers. comm.). After contact is made and prey defense is 
initiated, invertebrate predators may best respond to a prey’s defense 
by attacking the most vulnerable part of the prey. For example, aphids 
are more likely to be caught by predators when approached from the 
rear (Dixon 1958, Klingauf 1967). For caterpillar-shaped animals, the 
posterior may be the most vulnerable location also (sawfly larvae, Mor- 
row et al. 1976). Parasitoids that attack Baltimore checkerspot cater- 
pillars outside the communal webs maneuver carefully toward the hind 
end of the defensive hosts (Stamp 1982). Consequently, the proportion 


VOLUME 40, NUMBER 3 203 


of successful assaults may be greater on the posterior than near the 
head. 

In conclusion, when a caterpillar is smaller than the predator, the 
caterpillar is less likely to defend itself successfully. In this case, escape 
may be a more appropriate response. In contrast, when the body-size 
ratio of the caterpillar to the insect predator is large enough, a cater- 
pillar’s defense may be fairly effective. But caterpillars can defend 
only a limited space around themselves due to the particular stance 
they take. The maximal defendable space of caterpillars should change 
with body diameter, length of the moving (defending) portion of the 
body relative to total length, cuticular stretch, and physiological state. 
Insect enemies that cannot overwhelm a caterpillar may respond to 
the maximal defendable space by orienting to and attacking the more 
vulnerable (less defended) portion of the caterpillar’s body. 


ACKNOWLEDGMENTS 


I thank D. Bowers, S. Courtney and J. Myers for comments on the manuscript. I 
appreciate discussion on the topic with T. Friedlander, M. Hildebrand, J. Kingsolver, A. 
Shapiro and M. Tatar, and a reference provided by R. Lueschner. This research was 
done on a postdoctoral research fellowship at the University of California, Davis. 


LITERATURE CITED 


ALLEN, D. C., F. B. KNIGHT & J. L. FOLTz. 1970. Invertebrate predators of the jack- 
pine budworm, Choristoneura pinus, in Michigan. Ann. Entomol. Soc. Am. 63:59-64. 

BANKS, C. J. 1957. The behaviour of individual coccinellid larvae on plants. Brit. J. 
Anim. Behav. 5:12-24. 

BERISFORD, Y. C. & C. H. Tsao. 1975. Parasitism, predation, and disease in the bag- 
worm, Thyridopteryx ephemeraeformis (Haworth) (Lepidoptera: Psychidae). En- 
viron. Entomol. 4:549-554. 

BORNEMISSZA, G. F. 1966. An attempt to control ragwort in Australia with the cinnabar 
moth, Callimorpha jacobaeae (L.) (Arctiidae: Lepidoptera). Austral. J. Zool. 14:201- 
243. 

Brown, H. D. 1974. Defensive behavior of the wheat aphid, Schizaphis graminum 
(Rondani) (Hemiptera: Aphididae), against Coccinellidae. J. Entomol. (A) 48:157- 
165. 

CaTE, R. H., R. D. EIKENBERRY & R. D. MorRRISON. 1977. Preference for and effect 
of greenbug parasitism and feeding by Aphelinus asychis. Environ. Entomol. 6:547— 
500. 

COHEN, J. 1977. Statistical power analysis for the behavioral sciences. Rev. ed. Aca- 
demic Press, Orlando. 474 pp. 

DEMPSTER, J. P. 1971. The population ecology of the cinnabar moth Tyria jacobaeae 
L. (Lepidoptera, Arctiidae). Oecologia 7:26-67. 

DETHIER, V.G. 1959. Food-plant distribution and density and larval dispersal as factors 
affecting insect populations. Can. Entomol. 91:581-596. 

Dixon, A. F. G. 1958. The escape responses shown by certain aphids to the presence 
of the coccinellid Adalia decempunctata (L.). Trans. R. Entomol. Soc. (London) 110: 
319-334. 

1959. An experimental study of the searching behaviour of the predatory 

coccinellid beetle Adalia decempunctata (L.). J. Anim. Ecol. 28:259-281. 





204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1973. The biology of aphids. Edward Arnold, London. 58 pp. 

EISNER, T. & Y. C. MEINWALD. 1965. Defensive secretion of a caterpillar (Papilio). 
Science 150:1733. 

Evans, E. W. 1982. Feeding specialization in predatory insects: Hunting and attack 
behavior of two stinkbug species (Hemiptera: Pentatomidae). Am. Midl. Natur. 108: 
96-104. 

Evans, H. F. 1976. The role of the predator-prey size ratio in determining the effi- 
ciency of capture by Anthocoris nemorum and the escape reactions of its prey 
Acyrthosiphon pisum. Ecol. Entomol. 1:85—90. 

FELTWELL, J. 1982. Large white butterfly: The biology, biochemistry and physiology 
of Pieris brassicae (Linnaeus). Dr. W. Junk, The Hague. 535 pp. 

FITZGERALD, T. D. 1980. An analysis of daily foraging patterns of laboratory colonies 
of the eastern tent caterpillar, Malacosoma americanum (Lepidoptera: Lasiocam- 
pidae), recorded photoelectrically. Can. Entomol. 112:731-738. 

FLESCHNER, C. A. 1950. Studies on the searching capacity of the larvae of three 
predators of the citrus red mite. Hilgardia 20:233-265. 

FRANK, J. H. 1971. Carabidae (Coleoptera) as predators of the red-backed cutworm 
(Lepidoptera: Noctuidae) in central Alberta. Can. Entomol. 103:1089-1044. 

HEINRICH, B. 1979. Foraging strategies of caterpillars: Leaf damage and possible pred- 
ator avoidance strategies. Oecologia 42:325-337. 

HEPBURN, H. R. & H. LEvy. 1975. Mechanical properties of some larval cuticles. J. 
Entomol. Soc. S. Africa 38:131-140. 

Hicks, C. H. 1931. The hunt and capture of the prey of a digger wasp. Bull. S. Calif. 
Acad. Sci. 80:75-82. 

HUMPHRIES, D. A. & P. M. DRIVER. 1967. Erratic display as a device against predators. 
Science 156:1767-1768. 

Iwao, S. & W. G. WELLINGTON. 1970. The influence of behavioral differences among 
tent-caterpillar larvae on predation by a pentatomid bug. Can. J. Zool. 48:896—898. 

KLINGAUF, F. 1967. Abwehr- und Meideraktionen von Blattlausen (Aphididae) bei 
Bedrohung durch Raubern und Parasiten. Z. Angew. Entomol. 60:269-317. 

LANGSTON, R. L. 1957. A synopsis of hymenopterous parasites of Malacosoma in Cal- 
ifornia. Univ. Calif. Publ. Entomol. 14:1-50. 

Lopez, J. D., Jr., R. L. RiDGEWaAy & R. E. PINNELL. 1976. Comparative efficacy of 
four insect predators of the bollworm and tobacco budworm. Environ. Entomol. 5: 
1160-1164. 

MCFADDEN, M. W. 1968. Observations on feeding and movement of tobacco horn- 
worm larvae. J. Econ. Entomol. 61:352-356. 

MONTGOMERY, S. L. 1982. Biogeography of the moth genus Eupithecia in Oceania 
and the evolution of ambush predation in Hawaiian caterpillars (Lepidoptera: Geo- 
metridae). Entomol. Gen. 8:27-34. 

Morris, R. F. 1963. The effect of predator age and prey defense on the functional 
response of Podisus maculiventris Say to the density of Hyphantria cunea Drury. 
Can. Entomol. 95:1009-1020. 

1972. Predation by wasps, birds, and mammals on Hyphantria cunea. Can. 
Entomol. 104:1581-1591. 

Morrow, P. A., T. E. BELLAS & T. EISNER. 1976. Eucalyptus oils in the defensive oral 
discharge of Australian sawfly larvae (Hymenoptera: Pergidae). Oecologia 24:193- 
206. 

MYERS, J. H. & B. J. CAMPBELL. 1976. Predation by carpenter ants: A deterrent to the 
spread of cinnabar moth. J. Entomol. Soc. Brit. Col. 73:7-9. 

RABB, R. L. & F. R. LAwson. 1957. Some factors influencing the predation of Polistes 
wasps on the tobacco hornworm. J. Econ. Entomol. 50:778-784. 

RAUSHER, M. D. 1979. Egg recognition: Its advantage to a butterfly. Anim. Behav. 27: 
1034-1040. 

ROITBERG, B. D. & J. H. Myers. 1978. Adaptation of alarm pheromone responses of 
the pea aphid Acyrthosiphon pisum (Harris). Can. J. Zool. 56:103-108. 








VOLUME 40, NUMBER 3 205 


RUSSEL, R. J. 1972. Defensive responses of the aphid Drepanosiphum platanoides in 
encounters with the bug Anthocoris nemorum. Oikos 23:264-267. 

SCHAEFER, P. W. 1977. Attacking wasps, Polistes and Therion, penetrate silk nests of 
fall webworm. Environ. Entomol. 6:591. 

STAMP, N. E. 1982. Behavioral interactions of parasitoids and Baltimore checkerspot 
caterpillars (Euphydryas phaeton). Environ. Entomol. 11:100-104. 

STEINER, A. L. 1974. Unusual caterpillar-prey records and hunting behavior for a 
Podalonia digger wasp: Podalonia valida (Cresson). Pan-Pac. Entomol. 50:73-77. 

SToRCH, R. H. 1976. Prey detection by fourth stage Coccinella transversoguttata 
larvae (Coleoptera: Coccinellidae). Anim. Behav. 24:696-693. 

SUZUKI, N., Y. KUNIMI, S. UEMATSU & K. KOBAYASHI. 1980. Changes in spatial distri- 
bution pattern during the larval stage of the fall webworm, Hyphantria cunea Drury 
(Lepidoptera: Arctiidae). Res. Popul. Ecol. 22:273-283. 

SWYNNERTON, C. F. M. 1915. Experiments on some carnivorous insects, especially the 
driver ant Dorylus and with butterflies eggs as prey. Trans. Entomol. Soc. London 
1915, 317-350. 

TILMAN, D. 1978. Cherries, ants and tent caterpillars: Timing of nectar production in 
relation to susceptibility of caterpillars to ant predation. Ecology 59:686-692. 

WOLFGANG, W. J. & L. M. RIDDIFORD. 1981. Cuticular morphogenesis during contin- 
uous growth of the final instar larva of a moth. Tiss. Cell 13:757-772. 

Zak, J. H. 1974. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, New Jersey. 
620 pp. 


Received for publication 7 January 1986; accepted 8 April 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 206-213 


NATURAL HISTORY OF GNOPHAELA LATIPENNIS 
(BOISDUVAL) (ARCTIIDAE: PERICOPINAE) IN 
NORTHERN CALIFORNIA 


GEORGE L. GODFREY 


Illinois Natural History Survey, Natural Resources Building, 
607 E. Peabody Drive, Champaign, Illinois 61820 


AND 


LAURENCE CRABTREE 
P.O. Box 79, Chester, California 96020 


ABSTRACT. Diurnal Gnophaela latipennis (Boisduval) has a protracted summer 
flight and is univoltine with four larval instars, the second of which overwinters. Phe- 
nology and natural history of the egg, larva, pupa, and adult are discussed relative to 
the principal larval host, Hackelia californica (Gray) Johnston (Boraginaceae), in study 
areas in the northern Sierra Nevada. The egg, all larval instars, and pupa are described. 


Gnophaela latipennis (Boisduval) (Arctiidae: Pericopinae) (Figs. 1, 
2) is common to California, Oregon, and Washington, especially in the 
meadows and other forest openings of the Cascade, Sierra Nevada, and 
Coast ranges (Stretch 1872-1873, 1882, Dyar 1900, Powell & Hogue 
1979, T. D. Eichlin, pers. comm.). Stretch (1872-18783, 1882) noted its 
abundance in the Siskiyou Mountains and Sierra Nevada. Its range 
extends from Easton, Kittitas Co., Washington [U.S. National Museum 
record (D. C. Ferguson, pers. comm.)] to “southern California” (the 
Illinois Natural History Survey has a specimen labeled “Santa Monica, 
S. Cal., 1 May 1879’), but its current distribution does not appear to 
extend south of Kern Co. (Greenhorn Mountains at the southern end 
of the Sierra Nevada) (J. P. Donahue, pers. comm.). It has been col- 
lected as low as 244-305 m [Big Creek Nature Reserve (University of 
California), Monterey Co., California] but more often at elevations 
ranging from 396-1890 m (J. A. Powell, pers. comm.). 

Gnophaela latipennis occurs in localized populations (Stretch 1872- 
1873) as caterpillars (Figs. 5, 6) on boraginaceous hosts: Cynoglossum 
grande Douglas ex Lehmann, C. occidentale Gray, Hackelia califor- 
nica (Gray) Johnston, Mertensia sp., and Myosotis sp. (Stretch 1872- 
1873, 1882, Dyar 1900, Donahue 1979, Powell & Hogue 1979) and as 
conspicuous black and white, diurnal moths (Figs. 1, 2) that fly lazily 
during nectaring and oviposition. The flight behavior is similar to that 
of Gnophaela vermiculata (Grote) in the Rocky (Cockerell 1889) and 
Uintah mountains. 

Available literature on the natural history of G. latipennis is scant, 
and little is known about its phenology. The purpose of this paper is 


VOLUME 40, NUMBER 3 207 


\ f i 
an ae lel. side \ 
™ Tg Ee 


j. e « ~~ 
tie 1 Pe ie oe 





Fics. 1-6. Gnophaela latipennis. 1, adult male; 2, adult female; 3, undeveloped egg; 
4, eggs ready to hatch; 5, first-instar larva; 6, last-instar larva. 


to report on these matters relative to populations of G. latipennis in 
the northern reaches of the Sierra Nevada. 


STUDY SITES AND METHODS 


Field observations and collections focused on two areas during 1979- 
1983 in Plumas Co., California: Sunflower Flat, 1463 m, 4.8 km NWW 
of Chester; and Section 26 on Mud Creek Rim, 1676 m, 6.4 air km 


208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hackelia californica flowering period 












= o_4____e______e__e_____e—____#—__+_*+-® B 
(1981) x LEGEND 
(1 983) @ Field collected or observed 


D) Laboratory rearing 


O Extrapolated 
EGS eo Ae el lee ee ieee “fel 


or Hackelia not in bloom 





Adult . 2.5 «7)} _» 0c» tTpeeeft}_o — 0 . 


Pupa foo» #4. +e —___. 


Prepupa-?. 4 
Athulvistarsegtee ys arti 
ington ts 2 at Se eee oe ens oS PAUIS Cals 


retake. geet ee Seen Pee 2nd Instar 3, eee 


Ist Instar pai} eee 2 __o 2 


---+ — --e _e—___e—_ ee 0 0 00 __0 00 _0 o_o __0 __0 00 o—_e—___o__________»—_______#ee ----- #--- 
Larval Composite 











11] 15|19|23|27| 31! 4 | 8 |12|16|20 | 24 | 28| 2 | 6 | 10)14)|18 | 22|26|30| 3 | 7 | 11} 15) 19] 23| 27|31| 4 | 8 
13°17 21 25 2 | 2 6 10 14 18 22 26 30 4 8 12 16 20 24 july 5 9 13 17 21 25 | 2 6 10 
May June Seen July August September 


Fic. 7. Phenology of Gnophaela latipennis in relation to the flowering period of its 
principal host, Hackelia californica, near Chester, Plumas Co., California. 


NE of Chester. Both sites are typical Sierran montane forests disturbed 
by earlier timber harvesting activity. The dominantly coniferous vege- 
tation is a mixture of Abies concolor (Gordon & Glendinning) Lindley, 
Pinus lambertiana Douglas, P. jeffreyi Greville & Balfour, and Libo- 
cedrus (Calocedrus) decurrens (Torrey) Florin. The shrubby under- 
story is primarily Ceanothus velutinus Douglas ex Hook, Arctostaphy- 
los patula Greene, and Castanopsis sempervirens (Kellogg) Dudley. 
Hackelia californica is the most abundant herb, and Cynoglossum 
occidentale occurs infrequently. Additional observations were made in 
Butte Co., 11.2 air km SE (Brown’s Ravine, 1585 m) and 14.5 air km 
SE (1890 m) of Butte Meadows. These areas are 40 air km SW of 
Chester. 

Site examinations and observations began with receding snow cover 
in mid-May and continued with occasional unavoidable interruptions 
into September. Field data were composited into a generalized seasonal 
history (Fig. 7). Part of the generalization was extrapolated by deter- 
mining the durations of the egg and pupal stages under ambient lab- 
oratory temperature, and by noting the hatching of field-collected eggs 
to establish the onset of the first larval instar. 

Hinton’s (1946) setal nomenclature was used in the larval descrip- 
tions. 


VOLUME 40, NUMBER 3 209 


OBSERVATIONS AND DISCUSSION 


Adult. The adults (Figs. 1, 2) of Gnophaela latipennis, a univoltine 
species, exhibited a protracted summer flight period (Fig. 7). They 
emerged from field pupae by 27 June and were numerous and active 
during the first three weeks in July; but adults observed after that time 
appeared weak and ragged. The last field sighting of an adult during 
the 4-year period was 29 July. However, egg clusters were seen on 
three later occasions, one as late as 14 August. As the minimum hatch- 
time of a G. latipennis egg is about seven days (discussed under “Egg’’), 
the female that laid the eggs would have been alive at least through 7 
August. Thus, observed and extrapolated information indicates that 
adults fly between 27 June and 7 August in the Sierra Nevada study 
areas. In the Coast Range of California, adults have been observed as 
early as 25 May (8.0 air km SE Hayfork, Trinity Co.), 5 June (UC Big 
Creek Nature Reserve, 244-305 m, Monterey Co.), 8 June (emergence) 
(Buttercreek Meadows, 12.9 km W Hayfork, 1148 m, Trinity Co.), and 
9 June (Kelseyville, Lake Co.) (Powell, pers. comm.). 

Hackelia californica, the principal larval host at the study sites near 
Chester, was used extensively by adult Gnophaela latipennis for nec- 
tar. Its flowering period began approximately three weeks before the 
first G. latipennis adults emerged (heavy snow cover during the winter 
of 1982-1988 resulted in a late spring, hence the lack of blossoms on 
7 June 1983) and overlapped their flight period through 31 July (Fig. 
7). By the third week of July, however, most blossoms had been suc- 
ceeded by well-developed nutlets. These nutlets are covered with pric- 
kles (Munz 1970), and two adults were found stuck to them on 29 July 
and five on 23 July 1980. Gnophaela latipennis also nectared at the 
flowers of Cynoglossum grande (Butte Co.), C. occidentale, and He- 
lianthus sp. The latter, plus Senecio, were visited by Gnophaela ver- 
miculata in the Rocky Mountains (Cockerell 1889). 

Many pairs of G. latipennis were observed mating between 5-25 
July on nectar plants, larval host plants, adjacent vegetation, and once 
on the end of a Pinus jeffreyi branch. One mating pair remained 
coupled for 3 h; a second pair, for 10 h. 

Egg. Eggs occurred from 5-14 July (Fig. 7) and were usually found 
in clusters on the undersides of larval host plant leaves, as observed on 
Hackelia californica and Cynoglossum occidentale. However, eggs also 
were sporadically found on such unrelated plants as Pinus jeffreyi 
(Pinaceae) (22 eggs among needles), Stipa sp. (Gramineae), and Cea- 
nothus velutinus (Rhamnaceae) (10 eggs on the underside of a leaf). 

The number of eggs per cluster laid in the laboratory ranged from 
5-18 (x = 10.3/cluster, SD = 2.6, n = 7), whereas field-collected eggs 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


No. of Specimens 


30 






4th 


; : 2.36 [2.52 |2.68 
68 .84 1.00 1 28 1.44 1 60 2.28 2.44 2.60 


Head Capsule Width (mm) 


Fic. 8. Measurements, frequency, and distribution of Gnophaela latipennis head 
capsule widths based on field-collected and reared larvae. 


from larval host plants ranged from 1-21 (x = 10.0, SD = 6.5, n = 14). 
The egg clusters from Pinus jeffreyi, Stipa sp., and Ceanothus velu- 
tinus were disregarded in determining cluster size. The total egg com- 
plement of Gnophaela latipennis females was not fully determined, 
but a single female field-collected in copula on 5 July laid 157 eggs 
the following day, and a second female that emerged and mated in 
the laboratory produced 109 eggs. 

The newly laid egg (Fig. 3) is smooth, spherical, yellow, and mea- 
sured 1.25-1.82 mm in diameter (x = 1.31 mm, SD = 0.08, n = 6). 
Shortly before hatching, the head capsule and long, black, body setae 
were easily visible through the chorion (Fig. 4). The top of the egg 
collapsed near the head capsule. During hatching, the mandibles may 
be seen, under magnification, biting at the collapsed region to effect 
an exit hole for the larva. 

Hatching time was 7-8 days; in one laboratory situation, 19 of 34 
eggs hatched in seven days (15 failed to hatch) and in a second case, 
37 of 38 eggs hatched between seven and eight days (1 failed). The 
maximum stage duration of any field-collected eggs was seven days (2 
collected on 22 July 1983 hatched on 29 July). 

Larva. Four larval instars were determined by the distribution and 
frequency of head capsule widths of field-collected and reared larvae 
(Fig. 8): 

First instar (Fig. 5). Head capsule, width 0.60-0.68 mm (x = 0.62 


VOLUME 40, NUMBER 3 2 


Gap. 





Fics. 9-12. Gnophaela latipennis pupa. 9, habitus (dorsal); 10, cremaster (dorsal); 
11, head tubercles (dorsal); 12, in cocoon on Pinus jeffreyi (head towards branch). 


mm, SD = 0.02, n = 52), tan, setae translucent. Body translucent green- 
ish yellow with distinct black tubercles on T-2—Ab-9; cervical shield 
yellowish tan, setae translucent, majority of D and SD setae longer 
than body diameter and black; L and V setae shorter and translucent; 
D and SD on Ab-1-—Ab-8 unisetose; crochets in homoideous mesoseries. 
Active 25 July—25 August (Fig. 7). 

Second instar. Head capsule, width 0.84-1.00 mm (x = 0.89 mm, 
SD = 0.04, n = 34), reddish brown, setae tan to black. Body initially 
concolorous with first instar but cervical shield darkens, black stripes 
appear, SD and D1 tubercles become slightly iridescent bluish black; 
D and SD setae subequal in length to body diameter, most tubercles 
multisetose; crochets heteroideous mesoseries. Extends from 2 August— 
16 June (overwinters) (Fig. 7). 

Third instar. Head capsule, width 1.28—-1.68 mm (x = 1.49 mm, SD = 
0.10, n = 28), concolorous with preceding instar. Body yellow and black 
striped, tubercles iridescent blue; D and SD setae shorter than body 
diameter; otherwise similar to second instar. Seen from 22 May-—24 
June (Fig. 7). (The occurrence on 22 May indicates that some third 
instars may overwinter.) 


912 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fourth instar (Fig. 6). Head capsule, width 2.32-2.68 mm (x = 2.47 
mm, SD = 0.11, n = 15), reddish brown. Body concolorous with third 
instar. Seen 12 June-13 July; prepupae 21 June-12 July (Fig. 7). 

During the study period, only three of the five borages previously 
mentioned had G. latipennis larvae on them: Hackelia californica, 
Cynoglossum occidentale, and C. grande. The latter two species were 
less abundant and were considered minor hosts; C. grande did not 
occur in the study sites near Chester but was present in Butte Co. 
Hackelia californica seems to be an aggressive, early successional plant 
that becomes abundant in areas disturbed by logging activities. Several 
early instar larvae may feed on the same host-plant leaf, dispersing as 
they grow. Negative feeding results were obtained when two groups 
of five first-instar larvae were placed on Pinus jeffreyi and Ceanothus 
velutinus, two of the “nonhosts’’ on which eggs were found. 

The overwintering site of the second-instar larvae was not discov- 
ered. Larvae were observed feeding on Hackelia californica as late as 
1 September, but by 10 September, no larvae could be found on this 
perennial or under its basal leaves or elsewhere in known feeding areas. 
Neither could inactive larvae be found under H. californica in accu- 
mulated duff and around its woody root crowns in early spring. 

In 1982, spring larvae were detected feeding at Sunflower Flat on 
22 May; none had been seen there six days earlier. At Section 26, no 
larvae were found on 16 May when H. californica was 2.5 cm tall and 
snow patches still persisted, but by 30 May, larvae were seen feeding 
on this host plant when it was 7.6 cm tall. Of the three larvae collected 
on 22 May, two were third instars, which indicates that they also may 
overwinter; however, none were seen in the fall. 

The latest date that last-instar larvae were observed actively feeding 
(13 July) is one day later than the latest date for prepupae (12 July). 
The earliest date a prepupa was seen (21 June) coincides with the first 
pupal record (Fig. 7). Therefore, the prepupal time line in Fig. 7 
should be extended in both directions, but there are insufficient data 
to determine extention lengths. 

Pupa. Pupae occurred from 27 June-2 August (Fig. 7), and the 
duration of single pupae was 9-12 days (x = 10.5 days, n = 11) based 
on the pupation dates of field-collected, last-instar larvae, and the adult 
emergence dates. 

The pupa is black with conspicuous yellow to orange abdominal 
spots (Fig. 9), is slightly “hairy,” and “... suspended hammocklike 
...” (Powell & Hogue 1979) by its cremaster (Fig. 10) and two head 
tubercles (Fig. 11) inside a white, loose, lacelike cocoon (Fig. 12). Only 
two of the dozens of pupae (and three prepupae) found during this 
study were in Abies concolor foliage. Most were seen in young Pinus 


VOLUME 40, NUMBER 3 ONS 


jeffreyi at heights ranging from 1.8-3.7 m, where the cocoons had 
been spun solitarily among the needles. Much time was spent searching 
unsuccessfully for pupae in other tree and shrub species, but the ab- 
sence of pupae on other vegetation or on the ground does not preclude 
the possibility of undetected predation. For Gnophaela vermiculata, 
Bruce (1888) noted “... when full grown the whole brood [larvae] 
appears to make for the nearest detached rock where they spin their 
cocoons in angles and crevices, generally in clusters, and often covering 
each others [Sic] cocoons so thickly that many of the moths are not able 
to make their way through but die crippled.” Although one G. lati- 
pennis moth was seen in the field with deformed wings, obviously 
unable to escape from the cocoon, no general deformity problem was 
evident. 


ACKNOWLEDGMENTS 


We thank J. A. Powell, University of California (Berkeley), J. P. Donahue, Natural 
History Museum of Los Angeles County, T. D. Eichlin, California Department of Food 
and Agriculture, and D. C. Ferguson, Systematic Entomology Laboratory, U.S.D.A., for 
providing unpublished information; and D. J. Voegtlin for photographing Figs. 3-5. The 
constructive comments of Donahue, Voegtlin, J. K. Adams, J. K. Bouseman, and E. 
Levine were useful. We also thank M. L. Mini, Chester, California, for assisting in the 
field, and W. D. Dakan, Quincy, California, for identifying host plants. Several INHS 
staff members graciously aided us: L. LeMere, M. Hardin, G. A. Herendeen, and J. P. 
Sherrod (graphics); M. L. Williamson (word processing). Field observations and data 
collections were the efforts of the junior author whereas the senior author contributed 
the descriptions and interpretations. This paper was partially supported by Illinois Ag- 
ricultural Experiment Station Project 12-361 Biosystematics of Insects. 


LITERATURE CITED 


BRUCE, D. 1888. Description of mature larva of Gnophaela vermiculata, G. & R. 
Entomol. Am. 4:24. 

COCKERELL, T. D. A. 1889. The larva of Gnophaela vermiculata, G. & R. Entomol. 
Am. 5:57-58. 

DONAHUE, J. P. 1979. Strategies for survival—The cause of a caterpillar. Terra 17(4): 
3-9. 

Dyar, H. G. 1900. A parallel evolution in a certain larval character between Syntom- 
idae and the Pericopidae. Proc. Entomol. Soc. Wash. 4:407—409. 

HINTON, H. E. 1946. The homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Ent. Soc. 
London 97:1-37. 

Munz, P. A. 1970. A California flora. University of California Press, Berkeley. 1681 


pp. 

POWELL, J. A. & C. L. HOGUE. 1979. California insects. University of California Press, 
Berkeley. 388 pp. 

STRETCH, R. H. 1872-1878. Illustrations of the Zygaenidae & Bombycidae of North 
America. San Francisco. 242 pp. 

1882. Larva of Gnophaela hopfferi. Papilio 2:82-83. 





Received for publication 27 January 1986; accepted 14 May 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 214-217 


GROWTH OF THE BUCKMOTHS HEMILEUCA LUCINA AND 
H. MAIA (SATURNIIDAE) ON THEIR OWN AND 
ON EACH OTHER’S HOSTPLANTS 


NANcy E. STAMP 


Department of Biological Sciences, State University of New York, 
Binghamton, New York 13901 


AND 


M. DEANE BOWERS 


Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts 02138 


ABSTRACT. We compared larval growth of Hemileuca lucina and H. maia on their 
own and on each other’s hostplants. As expected, relative growth rate (RGR) of H. lucina 
was higher on its own host (Spiraea latifolia) than on the nonhost (Quercus prinoides). 
Unexpectedly, RGR of H. maia on its own host (Q. prinoides) and on the nonhost (S. 
latifolia) were low and similar. 


Two species of buckmoths, Hemileuca lucina Hy. Edw. and H. maia 
(Drury) (Saturniidae), occur in the northeastern United States. They 
overlap some in geographic range, but their hostplants are usually 
different. 

Hemileuca maia is widespread but uncommon in the eastern United 
States, occurring from Massachusetts and Michigan south to Texas and 
Florida, and west to Missouri and Illinois (Ferguson 1971). It is usually 
associated with scrub oak, Quercus ilicifolia Wang. (Fagaceae), al- 
though it has been reported feeding on other oak species (Q. laevus, 
Q. rubra, Q. stellata and Q. velutina) (Tietz 1972). It also has been 
found on willow (Salix sp.), poplar (Populus sp.) and cherry (Prunus 
sp.) (Tietz 1972). Although Tietz (1972) reports H. maia on Spiraea 
salicifolia, this may have been a mistaken identification. 

In comparison to H. maia, H. lucina occurs in Massachusetts, south- 
ern Maine and New Hampshire. Sometimes it is abundant, but popu- 
lations are quite local and subject to large fluctuations in density. The 
hostplant, Spiraea latifolia (Ait.) Borkh. (Rosaceae), is much more 
widespread than H. lucina. The reasons are unknown, but may be 
linked to thermal requirements of the larvae (Stamp & Bowers 1986a). 
Hemileuca lucina has been found occasionally on Betula populifolia, 
Prunus serotina and Quercus sp. (Tietz 1972, Bowers & Stamp 1986). 

The moths’ ranges overlap in some areas of Massachusetts. For ex- 
ample, a population of H. maia occurs in pine barrens at Montague, 
Mass. (D. Schweitzer, pers. comm.), and populations of H. lucina are 
scattered throughout Leverett, Mass., in wet fields with S. latifolia. 
These areas are all in Franklin Co. 


VOLUME 40, NUMBER 3 els, 


As with many hemileucines, these species overwinter as eggs (Tuskes 
1984). The larvae feed in the spring and pupate in the soil, where they 
aestivate until adult emergence in the fall. Phenologies in Massachu- 
setts are offset slightly, with egg hatch and adult flight later in H. 
maia. 

The objective of this study was to compare the abilities of H. lucina 
and H. maia to grow on their own and on each other’s hostplants. Oak 
leaves are relatively tough (Feeny 1970), especially compared to Spi- 
raea leaves, although leaf toughness increases in Spiraea with leaf age 
(Stamp & Bowers 1986b). The leaves of these species probably differ 
in other important ways as well, such as in water content, and presence 
and amount of tannins and other allelochemicals. Because the moths 
are congeneric and have similar life history traits (Ferguson 1971), we 
expected that they could eat each other’s hostplants but, because their 
food is different, that they would grow best on their own host species. 


METHODS 


For the experiment, H. lucina larvae were reared from an egg clus- 
ter collected on 14 April 1985 at Leverett, Mass., where the hostplant 
was S. latifolia. The egg cluster was kept in a refrigerator until 2 June; 
the larvae hatched on 6 June. Newly hatched H. maia larvae were 
collected from Quercus prinoides Willd. on 7 June 1985 at Barnard 
Valley (Nantucket), Mass. Caterpillars were kept in a growth chamber 
on a photoperiod of 16L:8D at 25°C during the day and 20°C at night, 
and were reared through the first instar on their own hostplants. At 
the beginning of the second instar, half of the larvae of each species 
were fed S. latifolia and half were fed Q. prinoides leaves. Larvae 
were given these diets throughout the second instar. 

At the beginning of the third instar, the growth tests were begun 
with larvae fed on the same diet they had during the second instar. 
For each treatment, 14 newly molted, unfed larvae were weighed and 
placed individually in Petri dishes, with a weighed sprig of S. latifolia 
or leaf of Q. prinoides. Sprigs of S. latifolia were used because indi- 
vidual leaves were small and dried quickly. We used unlignified sprigs 
(leaves young and stems still green) because new leaves were primarily 
what H. lucina ate in the field. The larvae were given average sized 
Q. prinoides leaves. Each dish had a piece of wet toweling taped to 
its top to maintain humidity. Larvae fed freely for 48 h. Then they 
were weighed again, frozen, dried at 50°C for 72 h and reweighed to 
obtain wet-weight-to-dry-weight conversion factors to estimate initial 
dry weight. To compare the response of larvae to the two hostplants, 
a standard index of growth, relative growth rate (RGR), was used 
(Waldbauer 1968, Slansky & Feeny 1977). 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Growth indices for third instar Hemileuca lucina and H. maia. Each 
treatment had 14 larvae. Means are indicated with +1 SE. Means with the same letter 
in that column are not significantly different at the 0.05 level using the Newman-Keuls 
multiple range test. 


Diet during second Mean dry weight per larva RGR Mean dry weight per larva 
instar and test period (mg) at beginning of test (mg/mg larva/day) (mg) at end of test 
H. lucina 
Spiraea ope OP2ra 0.45 + 0.03 a 13.9+ 10a 
Quercus 3.8 + 0.2b 0.34 + 0.01 b 7.8 +0.5b 
H. maia 
Quercus 46+ 0.2a 0.17 + 0.0lc 6.7 + 0.4b 
Spiraea 2.9 = O3c¢ 0.18 + 0.0lc 41+04c 


RESULTS AND DISCUSSION 


After rearing through the second instar, H. lucina larvae fed Spiraea 
were heavier than those fed Quercus, but were similar in estimated 
dry weight to H. maia fed Quercus (Table 1). As expected, RGR of 
H. lucina fed Spiraea was higher than for those fed Quercus. But, 
unexpectedly, RGRs for H. maia on Spiraea and Quercus were both 
low and similar. Because H. maia growth on Quercus was so slow, 
those larvae did not gain much weight during the test (Table 1). Con- 
sequently, H. lucina fed Spiraea weighed significantly more at the end 
of the test than larvae in the other treatments. 

The relative growth of H. lucina on Quercus was probably the same 
as that of third-instar H. lucina reared on old leaves of S. latifolia 
(0.34 + 0.01 SE on Quercus, and 0.35 + 0.01 SE on old Spiraea; two- 
sample t-test, df = 26, P > 0.50, 1 — 6 = 0.31 at a = 0.05; data for 
old Spiraea leaves from Stamp & Bowers 1986b). Thus, Quercus and 
old Spiraea leaves were similar as food for H. lucina, but it was not 
clear why. Hemileuca maia did not exhibit such a pattern; it grew 
equally slowly on both its hostplant Quercus and nonhost Spiraea. This 
suggests that H. maia simply grow slowly in comparison to H. lucina. 
Alternately, Q. prinoides may be a less suitable host than Q. ilicifolia, 
the usual host of H. maia. However, in northern New York, H. maia 
used both oak species, and in one area used QO. prinoides predominantly 
even though the other oak species was present (Cryan & Dirig 1977). 

The end result (in this case, middle of the third instar) was quite 
different dry weights among the treatments (Table 1). Hemileuca lu- 
cina fed on Quercus weighed but 56% of the weight attained by H. 
lucina reared on Spiraea. Even more striking, H. maia reared on its 
hostplant Quercus weighed only 49% of that attained by H. lucina on 
its hostplant Spiraea. Because adults of these two buckmoths are similar 


VOLUME 40, NUMBER 3 WANT 


in size (H. lucina somewhat smaller; Cryan & Dirig 1977), it seems 
likely that H. maia larvae may have a prolonged developmental period 
compared to that of H. lucina. Warrington (1985) found such an effect, 
with the combination of RGR and weight of prepupal larvae affecting 
the pupation date for four species of geometrids feeding on sycamore. 

Prolonged larval development in H. maia may have important ram- 
ifications for exposure to abiotic conditions, predators and parasites 
and, consequently, for the means by which H. maia cope with those 
factors, especially compared with H. lucina. 


ACKNOWLEDGMENTS 


We thank G. Puttick, F. Slansky Jr., and G. P. Waldbauer for comments on the 
manuscript, and G. Puttick for collecting H. maia and Q. prinoides for us. NES was 
supported by an award from the NYS-UUP Professional Development Committee. MDB 
was supported by the Clark Fund of Harvard University and NSF Grant BSR 8307353. 


LITERATURE CITED 


Bowers, M. D. & N. E. Stamp. 1986. Host plant exploitation by gregarious larvae: 
Effect of temperature and group size on buckmoths (Hemileuca lucina: Saturniidae). 
In review. 

CRYAN, J. F. & R. Diric. 1977. The moths of autumn: Buckmoths of the Pine Bush. 
Occ. Publ. No. 1. Pine Bush Historic Preservation Project, Albany, New York. 16 pp. 

FEENY, P. 1970. Seasonal changes in oak leaf tannins and nutrients as a cause of spring 
feeding by winter moth caterpillars. Ecology 15:565-581. 

FERGUSON, D. C. 1971. The moths of North America. Fascicle 20.2. Bombycoidea. E. 
W. Classey, Middlesex, England. 

SLANSKY, F., JR. & P. FEENy. 1977. Stabilization of the rate of nitrogen accumulation 
by larvae of the cabbage butterfly on wild and cultivated food plants. Ecol. Monogr. 
47:209-228. 

STAMP, N. E. & M. D. Bowers. 1986a. Hostplant exploitation by gregarious larvae: 
Thermal ecology of buckmoths (Hemileuca lucina: Saturniidae). In review. 

1986b. Hostplant exploitation by gregarious larvae: Quality and availability of 
food and its effect on buckmoths (Hemileuca lucina: Saturniidae). In review. 

TrETzZ, H. M. 1972. An index to the described life histories, early stages and hosts of 
the Macrolepidoptera of the continental United States and Canada. Vol. 1. A. C. 
Allyn, Sarasota, Florida. 536 pp. 

TuskEs, P. M. 1984. The biology and distribution of California Hemileucinae (Satur- 
niidae). J. Lepid. Soc. 38:281-309. 

WALDBAUER, G. P. 1968. The consumption and utilization of food by insects. Adv. 
Insect Physiol. 5:229-288. 

WARRINGTON, S. 1985. Consumption rates and utilization efficiencies of four species 
of polyphagous Lepidoptera feeding on sycamore leaves. Oecologia 67:460—463. 





Received for publication 7 February 1986; accepted 23 May 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 218-237 


THE SPECIES OF PSEUDEXENTERA (TORTRICIDAE) 


WILLIAM E.. MILLER 


Department of Entomology, University of Minnesota, 
St. Paul, Minnesota 55108 


_ ABSTRACT. Seventeen species of the nearctic genus Pseudexentera are recognized 
based on sometimes subtle differences in forewing pattern, and on one or more corrobora- 
tive differences in structure or larval host. Seventeen characters were examined, and 
study material consisted of more than 1200 pinned adults, 450 genital preparations, and 
500 wing preparations. Thirteen species occur only east of the Great Plains, three only 
westward, and one transcontinentally. Identities are revised for P. cressoniana (Clemens), 
the type species; also for P. faracana (Kearfott) and P. spoliana (Clemens). Pseudexen- 
tera caryana McDunnough proved to be a junior synonym of P. cressoniana. New species 
and their type localities are P. sepia (Cincinnati, Ohio), P. hodsoni (Oak Station, Pa.), P. 
knudsoni (Riviera Beach, Tex.), P. oreios (Rustler Park, Ariz.), and P. vaccinii (S. March, 
Ont.). Lectotypes are designated for P. bipustulana (Walker) (a junior synonym of P. 
costomaculana (Clemens)), and for P. oregonana (Walsingham). 


Pseudexentera is a nearctic genus of the tribe Eucosmini. The known 
larvae appear to be monophagous or stenophagous on woody plant 
foliage. The adults are among the earliest spring-flying insects. Life 
history data and adult capture records suggest that all species of Pseud- 
exentera are univoltine. At least three species are of economic interest: 
P. mali is a pest of apple, and P. spoliana and P. oregonana reach 
conspicuous numbers on oak and aspen, respectively. 

Grote (1877) proposed the new genus and species Exentera aprilana, 
and Heinrich (1928) characterized the genus by male genital and other 
characters without having examined the type specimen. When he dis- 
covered that E. aprilana is a species of Eucosma, Heinrich (1940) 
proposed the name Pseudexentera for the genus, designating Hedya 
cressoniana Clemens as type species, and indicating his earlier belief 
(Heinrich 1923) that H. cressoniana was a junior synonym of Sciaphila 
improbana Walker. McDunnough (1959) discovered that S. improbana 
is a species of Zeiraphera. He regarded Hedya cressoniana as the type 
species of Pseudexentera, but likewise misidentified it. Thus the genus 
does not now have a correctly identified type species. The case is being 
referred to the International Commission on Zoological Nomenclature 
as called for by Art. 70b of the International Code of Zoological No- 
menclature (8rd ed.). 

There are two main forewing patterns in the genus, excluding the 
striped form of some P. faracana (Fig. 2). The main patterns are 
exemplified by P. cressoniana (Fig. 1) and P. costomaculana (Fig. 22). 
When forewing pattern varies within species, it ranges between distinct 
and less distinct or diffuse. Although adults can be identified by exter- 
nal appearance once their variation is understood, the species in each 


VOLUME 40, NUMBER 3 219 


of the two main wing-pattern groups are confusingly similar. This 
similarity has thwarted taxonomic progress and fostered confusion, es- 
pecially with two of the earliest named species, P. cressoniana and P. 
spoliana, whose types remained incorrectly identified for more than a 
century. 


METHODS 


Pinned adults first were segregated by differences in forewing pat- 
tern, sometimes subtle ones. The resulting groups were then examined 
for differences in structure, larval host, adult phenology, and geog- 
raphy. The number of characters totaled 17. Structure refers to wing 
venation, genitalia, and body size. Either individual or statistical dif- 
ferences in structure were admissible. One distinguishing character 
state in structure or larval host was deemed sufficient to confirm spe- 
cific distinction. Such differences are not always explicitly cited in the 
text except for new species. If differences beyond forewing pattern 
were not found, the group in question was combined with the next 
most similar one. Final groupings thought to comprise species were 
then assigned names based on their similarity to types. 

The following structural characters and states or ranges proved use- 
ful in sorting and diagnosing the species. Both sexes: origin of forewing 
veins R, and R,;: separate (like M, and M, in fig. 2 of Heinrich 1923), 
approximate (like Rs and M, in fig. 1 of Heinrich), connate (like M; 
and CuA, in fig. 1 of Heinrich), stalked (like M, and CuA, in fig. 2 of 
Heinrich). Males: position of valval constriction: 4%, °%4, % distance be- 
tween valval base and apex; maximal valval length/maximal cucullus 
length ratio, the latter distance measured between upper and lower 
edges: 1.6 to 3.4; position of anal spine on cucullus: near middle, near 
lower edge; projections from lower edge of cucullus: absent, one small 
but conspicuous curved spine present, one or more inconspicuous 
spinelets or bumps present; radius of curvature of lower edge of cu- 
cullus: greater or less than cucullus height; apex of aedeagus: unmod- 
ified, falcate, snoutlike and thin, snoutlike and thick; length of fore- 
wing: 5.0 to 10.0 mm. Females: position of ostium bursae: starting 4, 
to 1% width of ostium bursae behind front edge of sternum 7; taper of 
forward end of sterigma: gradual if any, sharp; lightly sclerotized patch 
on corpus bursae: present, absent; size of signa: equal or subequal, 
unequal; symmetry of forward and rear halves of papillae anales: sub- 
symmetrical, asymmetrical; length <x width at maxima of one papilla 
analis: 0.08 to 0.16 mm’; length of forewing: 5.5 to 10.0 mm. 

Genitalia were prepared as described by Clarke (1941). Position of 
the ostium bursae was estimated by eye at 45x nominal magnification. 
Measurements of papillae anales are to the nearest 0.03 mm, and were 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


made from microprojections. Valval length/cucullus length ratios were 
also determined from microprojections. Some wing venation prepara- 
tions were permanent, but most were temporary. The latter were pre- 
pared by touching a capillary tube of xylol to a wing in a microscope 
field illuminated from below. Subsamples for venation study consisted 
of equal or nearly equal numbers of males and females when 20 or 
more adults of a species were available. When fewer adults were avail- 
able, all were used regardless of the sex ratio. Wing length includes 
tegula and fringe, and was measured without magnification to the 
nearest 0.5 mm. 

The letter n after a number denotes number of specimens under- 
lying a statement. A parenthetical number after a color name refers to 
a color in Smithe (1975). Colors were estimated in natural light enter- 
ing a window. Plant names follow Little (1979). 

I examined 1234 pinned adults, 479 genitalia preparations, and 544 
wing preparations from 20 sources. All nontypes bear the label “Vouch- 
er, W. E. Miller, 1986,” and those illustrated here are also marked. 
The sources, with abbreviations used in the text, are: AMNH, American 
Museum of Natural History, New York; ANSP, Academy of Natural 
Sciences of Philadelphia; BMNH, British Museum (Natural History), 
London; CNC, Canadian National Collections of Insects, Arachnids 
and Nematodes, Ottawa; Craig, W. S. Craig, Columbia, Mo.; CU, Cor- 
nell University, Ithaca, N.Y.; FMNH, Field Museum of Natural His- 
tory, Chicago; Heitzman, J. R. Heitzman, Independence, Mo.; INHS, 
Illinois Natural History Survey, Urbana; Knudson, E. C. Knudson, Bel- 
laire, Tex.; LACM, Natural History Museum of Los Angeles County, 
Los Angeles; Leuschner, Ronald Leuschner, Manhattan Beach, Cal.; 
Mather, Bryant Mather, Clinton, Miss.; MSUE, Michigan State Uni- 
versity Entomology Museum, East Lansing; NSM, Nova Scotia Museum, 
Halifax; UCB, University of California, Berkeley; UMMZ, University 
of Michigan Museum of Zoology, Ann Arbor; UMSP, University of 
Minnesota, St. Paul; USNM, National Museum of Natural History, 
Washington, D.C.; UWM, University of Wisconsin, Madison. 

Nomenclatural summaries consist mostly of primary works. Check 
lists are omitted except where they introduced new name combina- 
tions. Keys to species are omitted in the belief that matching specimens 
to diagnoses and illustrations is a suitable user alternative that saves 
space. 


Pseudexentera Heinrich (1940:243) 


Exentera (not Grote 1877:227); Heinrich (1923:172). 


Diagnosis. The genus has been characterized by Heinrich (1923) and Brown (1982). 
Both sexes: forewing lacking raised scale tufts, termen notched between veins CuA, and 


VOLUME 40, NUMBER 3 DOM 


M,. Males: lacking forewing costal fold; socii densely setose, fused basally, articulating 
with tegumen on a stem; valval neck partly clothed with dense spinelike setae, a tiny 
anal spine on cucullus. Females: sternum 7 deeply emarginate around ostium bursae; 
sterigma consisting of lamella postvaginalis; lamella antevaginalis absent; tergum 9 with 
two to four tiny backwardly projected spinelets, a trait not previously known; anterior 
apophyses longer than posterior apophyses; ductus bursae partly sclerotized in an elbow 
shape near middle; corpus bursae with two finlike signa. 

The genus appears to be monophyletic, the stemmed socii constituting a shared and 
perhaps derived character. 


Pseudexentera cressoniana (Clemens) 
(Figs. 1, 24, 41) 


Hedya cressoniana Clemens (1865:514) (lectotype: Virginia, no date, abdomen missing, 
forewing length 8.5 mm, Type ... 7222, designated by Darlington 1947, wings 
illustrated by Miller 1973a, in ANSP). 

Exentera improbana (not Walker 1863:337); Heinrich (1923:174) (part). 

Pseudexentera caryana McDunnough (1940:243) (holotype: female, St. David’s, Ont., 14 
April 1938, No. 5105, genitalia illustrated by McDunnough 1940, in CNC). NEW 
SYNONYMY. 


Diagnosis. Forewing pattern (Fig. 1) varying little between or within sexes, but females 
averaging slightly higher in contrast (90n). Forewing veins R, and R, usually (92%) 
stalked at origin, sometimes (8%) connate (48n). In males, valva constricted at % distance 
between base and apex, valval length/cucullus length ratio (spine included) 2.0 to 2.2, 
anal spine near lower edge of cucullus, aedeagus has a falcate apex (10n), a small curved 
spine projects from lower edge of cucullus (40n) (Fig. 24). The last is readily revealed 
without dissection by brushing away valval scales. In females, ostium bursae begins % to 
% its width behind front edge of sternum, forward end of sterigma tapers gradually if 
at all, corpus bursae spicule bases usually (94%) nowhere fused into a sclerotized patch, 
and signa unequal in size (Fig. 41) (18n). Forewing length of males 8.5 to 10.0 mm (40n), 
of females 8.0 to 10.0 mm (45n). 

Comments. This species was known most recently as P. caryana McDunnough. Hein- 
rich (1923) misidentified it, considering it to be a junior synonym of Sciaphila improbana 
Walker, which he also misidentified. When McDunnough (1959) discovered that the 
latter represented a species of Zeiraphera, he perpetuated Heinrich’s misidentification of 
P. cressoniana, apparently unaware that Darlington (1947) had designated a lectotype 
for the species. Although the lectotype lacks an abdomen, its well preserved forewing 
pattern (Miller 1973a: fig. 10) is diagnostic. The pattern of P. caryana matches it well. 
This species accounts for the fifth of six known misidentifications of Clemens olethreutine 
types (Miller 1973a, 1973b, 1974, 1979, 1985). 

I examined adults from Connecticut, Illinois, Iowa, Michigan, Mississippi, Missouri, 
New York, Pennsylvania, Texas, Virginia, and Wisconsin (AMNH, ANSP, Craig, Heitz- 
man, Knudson, Mather, MSUE, NSM, UCB, UMMZ, USNM, UWM). The study sample 
included the P. cressoniana lectotype and a P. caryana paratype. 

Biology. The larval host is Carya ovata (Mill.) K. Koch (McDunnough 1940). Adult 
capture dates range from 25 February to 16 May (90n). 


Pseudexentera faracana (Kearfott) 
(Figs. 2-4, 25, 42) 


Proteopteryx faracana Kearfott (1907:47) (lectotype male: Scranton, Pa., 21 April 1906, 
A. E. Lister, genit. prep. CH 15 Dec. 1919, designated by Heinrich 1923, forewing 
length 9.0 mm, in AMNH). 

Exentera faracana; Heinrich (1923:177). 

Pseudexentera faracana; Powell (1983:36). 

Eucosma haracana (not Kearfott 1907:46); Busck (1914:150). 


222 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Exentera spoliana (not Clemens 1865:513) (part); Heinrich (1923:175). 
Pseudexentera spoliana (not Clemens 1865:513); MacKay (1959:123), Brown (1982:595). 


Diagnosis. Forewing pattern of three slightly intergrading kinds, varying greatly be- 
tween and within sexes, females possibly averaging higher in contrast. Kinds of pattern 
are striped (Fig. 2) (7n), thick banded (Fig. 3) (8n), and thin banded (Fig. 4) (35n). 
Forewing veins R, and R,; usually (73%) approximate or connate at origin, sometimes 
(27%) stalked or separate (52n). In males, valva constricted at % distance between base 
and apex, valval length/cucullus length ratio 2.3 to 2.7, anal spine near lower edge of 
cucullus, radius of curvature of lower edge of cucullus less than cucullus height, lower 
edge of cucullus may have up to four inconspicuous projections ranging in shape from 
bumps to spinelets, aedeagus has thin snoutlike apex (Fig. 25) (16n). In females, ostium 
bursae begins % to % its width behind front edge of sternum, forward end of sterigma 
tapers gradually if at all, corpus bursae spicule bases fused on one side near ductus bursae 
into a lightly sclerotized patch, signa unequal in size (Fig. 42) (25n). Forewing length of 
males 7.5 to 9.0 mm (19n), of females 7.0 to 8.5 mm (31n). 

Comments. This species was most recently known partly as P. spoliana and partly as 
P. faracana. The former reflects Heinrich’s (1923) misidentification of P. spoliana, which 
has a forewing pattern different from the three P. faracana patterns. Kearfott (1907) 
and Heinrich (1923) mused that one species with variable forewing pattern might be 
involved. Heinrich kept them separate because there were so few typelike examples of 
P. faracana at the time, the lectotype having the striped pattern. The three patterns here 
might indeed represent more than one species, but the information available now does 
not permit more than one to be recognized. Larval hosts are unknown for the striped 
and thick-banded adults. The larva of the thin-banded adult was described by MacKay 
(1959). 

I examined adults from Connecticut, Illinois, Michigan, Mississippi, Missouri, New 
York, Ohio, Pennsylvania, Virginia, West Virginia, and Wisconsin (AMNH, ANSP, CNC, 
CU, FMNH, Heitzman, INHS, LACM, Mather, MSUE, UCB, USNM, UWM). The study 
sample included the P. faracana lectotype and a paralectotype, the pinned adult and 
genit. prep. No. 252 of Brown (1982), and thin-banded adults reared from Castanea 
dentata (Marsh.) Borkh. (Hopk. U. S. 1134) (4n). 

Biology. The larva rolls the leaves of Castanea dentata, American chestnut, and the 
pupa overwinters on the ground (Busck 1914, MacKay 1959). There is one generation 
per year. Adult capture dates range from 25 February to 10 May (45n). 

Although American chestnut was nearly exterminated by the blight fungus Endothia 
parasitica (Murr.) Anders. and Anders. early in this century, the host persists as stump 
sprouts and occasional disfigured trees. These host remnants could explain how % of the 
study sample originated since 1950, two adults as recently as 1982. I captured an adult 
at stump sprouts. Alternate Castanea hosts could also be involved. 


Pseudexentera sepia Miller, new species 


(Figs. 5, 26, 43) 
Exentera spoliana (not Clemens 1865:513) (part); Heinrich (1923:175). 


Diagnosis and description. Forewing pattern (Fig. 5) varying little between or within 
sexes, females possibly averaging higher in constrast (22n). Forewing veins R, and R, 
usually (81%) approximate or connate at origin, sometimes (19%) separate or stalked 
(21n). In males, valva constricted at % distance between base and apex, valval length/ 
cucullus length ratio 2.2 to 2.6, anal spine near lower edge of cucullus, lower edge of 
cucullus has one or more inconspicuous projections ranging in shape from bumps to 
spinelets, aedeagus has thin snoutlike apex (Fig. 26) (7n). In females, ostium bursae begins 
% to % its width behind front edge of sternum, forward end of sterigma tapers gradually 
if at all, corpus bursae spicule bases fused on one side near ductus bursae into a lightly 
sclerotized patch, signa unequal in size (Fig. 43) (11n). Forewing length of males 7.0 to 
8.5 mm (8n), of females 7.0 to 8.5 mm (14n) (holotype 7.5 mm). Head. Labial palpus 
mixed white and brown, second segment subequal in length to eye diameter, apical 


VOLUME 40, NUMBER 3 223 


segment 4 length of second segment; front and crown mixed white and brown. Thorax. 
Mixed white and brown dorsally, shiny white ventrally; front and middle legs mixed 
white and brown, hind legs paler, tarsi of all legs banded; forewing upper-side dark 
markings near burnt umber (22) and raw umber (23), underside pale brown; hindwing 
upper side pale brown, underside paler. Abdomen. Mixed white and brown dorsally, 
paler ventrally. Male Genitalia. Vesica with 14 to 29 deciduous cornuti (5n). 

Type data. Holotype female: Cincinnati, OHIO, A. F. Braun, 7 April 1906, genit. 
prep. DH 813813 (AMNH). Nineteen paratypes (AMNH, CNC, LACM, UCB, UMSP, 
INHS, Mather, Heitzman, USNM): ILLINOIS: Putnam Co., 31 March 1940, M. O. Glenn, 
2 genit. prep. DH 630812 (Fig. 43); MISSISSIPPI: Vicksburg, 19 Feb. 1982, B. Mather, 
2 genit. prep. WEM 510841; MISSOURI: Kansas City, 19 April 1971, J. R. Heitzman, 2 
genit. prep. SMG 905823; same data except 6 genit. prep. SMG 907824 (this and preced- 
ing captured in copula); same data except 18 April 1973 (Fig. 5), wing prep. MGP 24, 
2 genit. prep. WEM 107844; same data except 18 April, 6 genit. prep. SMG 905822 (Fig. 
26); Grundy Co., 5 April 1980, J. R. Heitzman, ¢ genit. prep. SMG 1108828; same data 
except 2 genit. prep. SMG 1108827; PENNSYLVANIA: New Brighton, 16 March 1903, 
H. D. Merrick, 6, Proteopteryx spoliana Clem. AB 1920; same data except 6 genit. prep. 
CH 3, 30 Jan. 1920, Exentera spoliana Clem. var.; same data except 18 March 1903, 4, 
Exentera spoliana Clem. AB; same data except 30 April 1907, 9; Pittsburgh, 22 April 
1906, H. Engel, 2 genit. prep. WEM 175854; QUEBEC: Ottawa Co., 1-7 April, 2° genit. 
prep. LKM 824761; Old Chelsea, 30 April 1937, T. N. Freeman, 2 genit. prep. WEM 
107843; same data except 25 April 1935, W. J. Brown, wing prep. MGP 20, 6 genit. prep. 
Exen 4A; Aylmer, 9 May 1932, G. S. Walley, 2 genit. prep. Exen 4A, wing prep. MGP 
19; WISCONSIN: Milwaukee, 25 April 1915, H. M. Bower, 2 genit. prep. WEM 175853; 
Oneida Co., 20 May 1961, H. M. Bower, ¢ genit. prep. PB 221. 

Comments. Heinrich (1923) included this species in what is here considered P. fara- 
cana, sometimes adding “var.” to determinations. The new species most resembles thin- 
banded P. faracana. Among departures in forewing pattern, the basal patch is obscure, 
unlike that of P. faracana (Figs. 4, 5). Also, it is a smaller-bodied insect, based on forewing 
length. Forewing length of females, the more abundant sex in the study sample, averages 
7.6 mm (l11n) compared to 8.2 mm (19n) in P. faracana. The difference, 0.6 mm, is 
statistically significant (t = 3.7, Pa < 0.001). Although this difference represents only 
7% of the longer forewing, it denotes a 25% lighter body weight because of the expo- 
nential relation between these variables (Miller 1977). 

The species name refers to the family of hues marking the forewing. 

Biology. The larval host is unknown. Adult capture dates range from 19 February to 
20 May (21n). 


Pseudexentera hodsoni Miller, new species 


(Figs. 6, 27, 44) 


Exentera spoliana (not Clemens 1865:513) (part); Heinrich (1923:175). 


Diagnosis and description. Forewing pattern (Fig. 6) varying little between or within 
sexes (36n). Forewing veins R, and R, usually (97%) approximate or connate at origin, 
sometimes (3%) separate (34n). In males, valva constricted at % distance between base 
and apex, valval length/cucullus length ratio 2.7 to 3.4, anal spine near lower edge of 
cucullus, radius of curvature of lower edge of cucullus exceeds cucullus height, lower 
edge of cucullus may have up to two inconspicuous projections ranging in shape from 
bumps to spinelets, and aedeagus has thin snoutlike apex (Fig. 27) (18n). In females, 
ostium bursae begins % to 4 its width behind front edge of sternum, forward end of 
sterigma tapers gradually if at all, corpus bursae spicule bases fused on one side near 
ductus bursae into a lightly sclerotized patch, signa unequal or subequal in size (Fig. 44) 
(13n). Forewing length of males 7.0 to 8.5 mm (19n) (holotype 8.0 mm), of females 7.5 
to 9.0 mm (17n). Head. As described for P. sepia. Thorax. As described for P. sepia 
except forewing upper-side dark markings near dusky brown (19), and hindwing upper 


994 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


side grayish brown. Abdomen. As described for P. sepia. Male Genitalia. Vesica con- 
taining 22 to 34 deciduous cornuti (4n). 

Type data. Holotype male: Oak Station, Allegheny Co., PENNSYLVANIA, 10 April 
1910, F. Marloff, genit. prep. CH 20, 13 Feb. 1920, Exentera spoliana Clem. det. C. H. 
(USNM). Thirty-five paratypes (AMNH, Craig, CU, Heitzman, INHS, Knudson, Mather, 
USNM, UCB): FLORIDA: Pensacola, 7 Feb. 1962, S. Hills, 6 genit. prep. JAP 1424; 
ILLINOIS: Putnam Co., 7 April 1963, M. O. Glenn, 6; same data except 13 April 1941, 
8 genit. prep. WEM 168851; LOUISIANA: Baton Rouge, 14 Feb. 1970, G. Strickland, 9; 
MISSISSIPPI: Miss. State Univ., 1 March 1976, C. T. Bryson, 6 genit. prep. SMG 916822; 
same data except 19 March, 2 genit. prep. SMG 1029821 (Fig. 44); Hinds Co., 31 Jan. 
1967, B. Mather, 6 genit. prep. JAP 2441; same data except 13 Feb. 1959, 6; same data; 
same data except 6 genit. prep. SMG 916823; same data except 10 March 1963, 6 genit. 
prep. JAP 1578; same data except 16 March 1963, 2 genit. prep. JAP 1719; Rankin Co., 
3 April 1960, B. Mather, 2 genit. prep. JCL 1107836; Bovina, Warren Co., 7 March 1975, 
B. Mather, 2 genit. prep. SMG 915824; same data except 2 genit. prep. SMG 916821; 
same data except 14 March 1972, 2 genit. prep. JCL 1128831; Vicksburg, Warren Co., 
4 March 1980, B. Mather, 2; same data except 6; same data except 138 March 1981; same 
data except 30 March 1979, 2 genit. prep. WEM 207841; Gulfport, 31 March 1978, R. 
Kergosien, 2 genit. prep. SMG 916834; MISSOURI: “C. Mo.,” reared from red oak, C. 
V. Riley, é genit. prep. LKM 824765, Proteopteryx spoliana Clem. det. Walsingham 
1887; Jackson Co., 13 April 1972, J. R. Heitzman, ¢ genit. prep. WEM 207844; same 
data except 18 April 1979, 6 genit. prep. WEM 241852; Benton Co., 17 April 1970, J. R. 
Heitzman, 6; St. Genevieve Co., 10 April 1981, J. R. Heitzman, 2 genit. prep. WEM 
304851; Columbia, 10 April 1971, W. S. Craig, 2 genit. prep. JCL 1118833 (Fig. 6); same 
data except 12 April 1972, 2 genit. prep. JCL 1118834; same data except 16 April 1971, 
2; same data except 18 April, 6 genit. prep. WEM 207843; same data except 24 April, 9; 
Kirkwood, reared from “L. oak,” 1908, Murtfeldt, 2 genit. prep. WEM 197844; PENN- 
SYLVANIA: Allegheny Co., 30 April 1911, F. Marloff, 2 genit. prep. JCL 11178310; 
TEXAS: Anderson Co., 15 March 1983, E. C. Knudson, ¢ genit. prep. ECK 584; Harris 
Co., 5 Feb. 1984, E. C. Knudson, 6 genit. prep. ECK 817 (Fig. 27); same data except 19 
Feb. 1985, 4. 

Comments. Heinrich (1923) included this species in what is here considered Pseudex- 
entera faracana. The new species superficially resembles thin-banded P. faracana but 
differs structurally. The mate valval constriction is at % the distance between valval base 
and apex, compared to % in P. faracana; and the lower edge of the male cucullus is 
slightly rounded, its curvature forming a circle whose radius exceeds cucullus height, 
while the lower edge of the cucullus in P. faracana is greatly rounded, its curvature 
forming a circle whose radius does not exceed cucullus height (Figs. 25, 27). 

The species is named for Alexander C. Hodson, distinguished entomologist, teacher, 
and administrator. 

Biology. The larval host is Quercus (2n), the species being given on labels as “L. oak” 
and “red oak.” According to Riley’s label information, the larva “slightly folds . . . leaves 
in May, pupates in (a) tough cocoon on (the) ground, and emerges early the following 
spring.” Adult capture dates range from 31 January to 30 April (34n). 


Pseudexentera knudsoni Miller, new species 


(Figs. 7, 28, 45) 


Diagnosis and description. Forewing pattern (Fig. 7) varying little within or between 
sexes (8n). Forewing veins R, and R, connate, approximate, or stalked at origin (8n). In 
males, valva constricted at % distance between base and apex, valval length/cucullus 
length ratio 2.4 to 2.7, anal spine near lower edge of cucullus, lower edge of cucullus 
may have up to two inconspicuous projections ranging in shape from bumps to spinelets, 
aedeagus has falcate apex (Fig. 28) (5n). In females, ostium bursae begins % its width 


VOLUME 40, NUMBER 3 DOR 


behind front edge of sternum, forward end of sterigma tapers gradually if at all, corpus 
bursae spicule bases fused on one side near ductus bursae into a lightly sclerotized patch, 
and signa unequal in size (Fig. 45) (2n). Forewing length of males 6.5 to 7.5 mm (6n) 
(holotype 7.0 mm), of females 7.0 to 7.5 mm (2n). Head. As described for P. sepia. 
Thorax. Mixed white, brown, and sometimes orange dorsally, shiny white ventrally; front 
and middle legs mixed white and brown, hind legs paler, tarsi of all legs banded; forewing 
upper-side dark markings near olive-brown (28) and glaucous (79); hindwing upper side 
pale brown, underside paler. Abdomen. As described for P. sepia. Male Genitalia: Vesica 
containing 19 to 29 deciduous cornuti (5n). 

Type data. Holotype male (Fig. 7): Riviera Beach, Kleberg Co., TEXAS, 24 Feb. 1984, 
E. C. Knudson, genit. prep. WEM 168855 (USNM). Seven paratypes (Knudson, USNM, 
UMSP): TEXAS: San Antonio, 2 April 1978, 4; same data except 6 May, 6 genit. prep. 
WEM 154852; Canyon Lake, Comal Co., 8 May 1982, 6 genit. prep. ECK 325 (Fig. 28); 
Conroe, Montgomery Co., 9 March 1982, 6 genit. prep. ECK 570; Benbrook, Tarrant Co., 
6 genit. prep. WEM 168856; Austin, 13 April 1979, ° genit. prep. WEM 15852; Sam 
Houston Nat. For., San Jacinto Co., 29 March 1978, 2 genit. prep. WEM 35852 (Fig. 45); 
all E. C. Knudson. 

Comments. This species most resembles thin-banded P. faracana. Its aedeagus has a 
faleate apex, whereas that of P. faracana has a thin snoutlike apex (Figs. 25, 28). Also, 
it is a smaller-bodied insect, based on forewing length. Forewing length of males, the 
more abundant sex in the study sample, averages 6.9 mm (6n) compared to 8.4 mm 
(18n) in P. faracana. The 1.5 mm difference is statistically significant (tf = 7.2, Pa < 
0.001). Although this difference represents only 18% of the larger P. faracana forewing 
length, it denotes a 50% body weight difference because of the exponential relation 
between these variables (Miller 1977). 

The species is named for Edward C. Knudson, its discoverer. 

Biology. The larval host is unknown. Adult capture dates range from 24 February to 
8 May (8n). 


Pseudexentera haracana (Kearfott) 
(Figs. 8, 29, 46) 


Proteopteryx haracana Kearfott (1907:46) (lectotype female: Hunter’s Range, Pike Co., 
Pa., 1 May 1906, selected by C. Heinrich, designated by Klots 1942, forewing length 
6.5 mm, in AMNH). 

Exentera haracana; Heinrich (1923:176). 

Pseudexentera haracana; Powell (1983:36). 


Diagnosis. Forewing pattern (Fig. 8) varying little between or within sexes (67n). 
Forewing veins R, and R, usually (82%) connate or approximate at origin, sometimes 
(18%) stalked or separate (33n). In males, valva constricted at % distance between base 
and apex, valval length/cucullus length ratio 2.3 to 2.7, anal spine near lower edge of 
cucullus, lower edge of cucullus has up to four inconspicuous projections ranging in shape 
from bumps to spinelets, and aedeagus has thick snoutlike apex (Fig. 29) (14n). In fe- 
males, ostium bursae begins %, to % its width behind front edge of sternum, forward 
end of sterigma tapers gradually if at all, corpus bursae spicule bases fused on one side 
near ductus bursae into a lightly sclerotized patch, signa unequal in size (Fig. 46) (16n). 
Forewing length of males 6.0 to 8.5 mm (32n), of females 6.0 to 8.0 mm (85n). 

Comments. J examined adults from Connecticut, Florida, Michigan, Minnesota, Mis- 
sissippi, Missouri, New Jersey, New York, Nova Scotia, Pennsylvania, Texas, and Wis- 
consin (AMNH, ANSP, Craig, Heitzman, Knudson, LACM, Mather, MSUE, NSM, UCB, 
UMSP, UWM). The study sample included paralectotypes (2n) and the lectotype. Adults 
superficially resemble Gretchena delicatana Heinrich; the two species are often mixed 
in collections. 

Biology. The larval host is unknown. Adult capture dates range from 26 February to 
15 June (67n). 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pseudexentera oreios Miller, new species 


(Figs. 9, 30, 47) 


Diagnosis and description. Forewing pattern (Fig. 9) varying little between or within 
sexes (7n). Forewing veins R, and R,; approximate at origin (7n). In males, valva con- 
stricted at %4 distance between base and apex, valval length/cucullus length ratio 2.8 to 
2.9, anal spine near lower edge of cucullus, lower edge of cucullus may have one or 
more inconspicuous projections ranging in shape from bumps to spinelets, aedeagus has 
thin snoutlike apex (Fig. 30) (4n). In females, ostium bursae begins %, to % its width 
behind front edge of sternum, forward end of sterigma tapers gradually, corpus bursae 
spicule bases fused on one side near ductus bursae into a lightly sclerotized patch, signa 
subequal in size (Fig. 47) (3n). Forewing length of males 7.5 to 8.0 mm (4n) (holotype 
8.0 mm), of females 7.0 to 8.0 mm (3n). Head. As described for P. sepia except for white 
front and orange crown. Thorax. Similar to crown hues dorsally, shiny white ventrally; 
legs as described for P. sepia; forewing upper-side dark markings near tawny (38) and 
raw umber (223), the white and silver ocellus conspicuous, underside brown; hindwing 
upper- and undersides brown. Abdomen. Brownish white dorsally, paler ventrally. Male 
Genitalia. Vesica with 25 to 32 deciduous cornuti (3n). 

Type data. Holotype male (Fig. 9): Rustler Park, Chiricahua Mts., ARIZONA, 3 July 
1972, 8500’ (2600 m), J. Powell, genit. prep. WEM 304853 (UCB). Six paratypes (UCB, 
UMSP): ARIZONA: Madera Canyon, Santa Rita Mts., Santa Cruz Co., 1-3 Aug. 1970, P. 
Rude, 6 genit. prep. WEM 65853; same data except 6 genit. prep. WEM 307851 (Fig. 
30); same data except 6 genit. prep. JAP 3614; same data except 2 genit. prep. WEM 
307853; Parker Canyon Lk., Cochise Co., 19 July 1972, J. Powell, 2 genit. prep. WEM 
35853 (Fig. 47); same data except 2 genit. prep. WEM 307854. 

Comments. This species most resembles P. haracana but differs structurally. The male 
valval constriction is at % the distance between valval base and apex, compared to % in 
P. haracana; and the snoutlike aedeagal apex is thin compared to that in P. haracana. 

The species name denotes mountain dwelling. 

Biology. The larval host is unknown. Adult capture dates range from 3 July to 3 
August (7n). 


Pseudexentera spoliana (Clemens) 
(Figs. 10, 11, 31, 48) 


Hedya spoliana Clemens (1865:513) (lectotype female: Virginia, no date, designated and 
illustrated by Miller 1973a, forewing length 8.0 mm, only right wings remaining, in 
ANSP). 

Exentera improbana (not Walker 1863:337); Heinrich (1923:174). 

Pseudexentera improbana (not Walker 1863:337); Heinrich (1940:242) (part), Mc- 
Dunnough (1940:244), Freeman (1942:213). 

Pseudexentera cressoniana (not Clemens 1865:514); McDunnough (1959:2). 

Pseudexentera spoliana; Miller (1973a:223). 


Diagnosis. Forewing pattern varying within sexes, overlapping between sexes, females 
averaging higher in constrast (Figs. 10, 11) (409n). Forewing veins R, and R, usually 
(83%) connate or stalked at origin, sometimes (17%) approximate (52n). In males, valva 
constricted approximately at middle, valval length/cucullus length ratio 1.8 to 2.1, anal 
spine near lower edge of cucullus, lower edge of cucullus lacks projections, aedeagus has 
falcate apex (Fig. 31) (44n). In females, ostium bursae begins % to 1% its width behind 
front edge of sternum, forward end of sterigma tapers gradually if at all, corpus bursae 
spicule bases usually (98%) nowhere fused into a sclerotized patch, signa subequal in size 
(Fig. 48) (62n). Forewing length of males 7.0 to 9.5 mm (194n), of females 6.5 to 9.0 
mm (217n). 

Comments. The nomenclatural history of this species intertwines with that of P. cres- 


VOLUME 40, NUMBER 3 Za 


soniana. Pseudexentera spoliana was long known as P. improbana, and most recently 
as P. cressoniana (McDunnough 1959). Because the syntype status of an earlier lectotype 
(Darlington 1947) seemed doubtful, a more plausible lectotype was designated (Miller 
1973a). This action was not destabilizing because the specimens involved are conspecific. 
Although the lectotype lacks an abdomen, its well preserved forewing pattern (Miller 
1973a: fig. 45) is diagnostic. This species accounts for the last of six known misidentifi- 
cations of Clemens olethreutine types (Miller 1973a, 1973b, 1974, 1979, 1985, and earlier 
in this paper). 

I examined adults from Colorado, Connecticut, District of Columbia, Illinois, Indiana, 
Iowa, Louisiana, Massachusetts, Michigan, Mississippi, Missouri, New Brunswick, New 
Hampshire, New Jersey, New York, Nova Scotia, Ohio, Ontario, Pennsylvania, Quebec, 
Texas, Virginia, and Wisconsin (AMNH, ANSP, CNC, Craig, CU, FMNH, Heitzman, 
INHS, Knudson, LACM, Mather, MSUE, NSM, UCB, UMMZ, UMSP, UWM). The study 
sample included the lectotype, adults whose wings were illustrated by Freeman (1942) 
(3n), and adults reared from Quercus rubra L. (24n) and Q. sp. (17n). The larva was 
described by MacKay (1959:122) 

Biology. The larva feeds on Quercus rubra and perhaps other oaks, rolling the leaves. 
There is one generation per year (Heinrich 1923, Freeman 1942). Adult capture dates 
range from 11 February to 30 May (363n). 


Pseudexentera mali Freeman 
(Figs. 12, 13, 32, 49) 


Pseudexentera mali Freeman (1942:218) (holotype male: Bell’s Corners, Ont., reared 
from Malus sylvestris (L.) Mill., 7 Feb. 1942, J. McDunnough, wings illustrated by 
Freeman 1942, No. 5384 in CNC), Chapman & Lienk (1971:52). 


Diagnosis. Forewing pattern varying within sexes, overlapping between sexes, females 
averaging higher in contrast (Figs. 12, 13) (73n). Forewing veins R, and R; usually (83%) 
stalked or connate at origin, sometimes (17%) approximate (41n). In males, valva con- 
stricted approximately at middle, valval length/cucullus length ratio 1.7 to 1.9, anal 
spine near lower edge of cucullus, lower edge of cucullus lacks projections, aedeagus has 
faleate apex (Fig. 32) (11n). In females, ostium bursae begins % to 1% its width behind 
front edge of sternum, forward end of sterigma tapers gradually if at all, corpus bursae 
spicule bases nowhere fused into a sclerotized patch, signa unequal or subequal in size 
(Fig. 49) (16n). Forewing length of males 6.5 to 8.0 mm (38n), of females 6.0 to 7.5 mm 
(39n). 

Comments. [ examined specimens from Michigan, Missouri, New York, Nova Scotia, 
Ontario, Quebec, and Wisconsin (Craig, CNC, CU, Heitzman, MSUE, NSM, UCB, USNM, 
UWM). The study sample included paratypes (3n), holotype, and adults reared from 
Malus sylvestris (14n). 

Biology. The larva feeds on Malus spp., mining buds and folding leaves. There is one 
generation per year, and the pupa winters on the ground (Chapman & Lienk 1971). 
Adult capture dates range from 13 March to 6 June (58n). The species is considered a 
pest of apple. 


Pseudexentera oregonana (Walsingham) 
(Figs. 14, 15, 33, 50) 


Semasia oregonana Walsingham (1879:62) (lectotype male: “Camp Watson, Grant Co., 
Ore., March-April 1872 ... Type ...,” genit. prep. No. 5724, selected by N. S. 
Obraztsov, designated here, forewing length 9.0 mm, in BMNH). 

Exentera improbana oregonana; Heinrich (1923:175). 

Pseudexentera oregonana; McDunnough (1940:244), Freeman (1942:213), MacKay (1959: 
122, 1962:640, 1965:668). 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-23. Pseudexentera wings, some images reversed. (1) cressoniana 6, Colum- 
bia, Mo.; (2) faracana 6, Putnam Co., Ill., striped; (8) faracana °, New Brighton, Pa., 
thick banded; (4) faracana 2, Falls Church, Va., thin banded; (5) sepia 2, Kansas City, 
Mo.; (6) hodsoni 2, Columbia, Mo.; (7) knudsoni holotype 6; (8) haracana 2, New Lisbon, 
N. J.; (9) oreios holotype 6; (10) spoliana 6, Independence, Mo., diffuse pattern; (11) 
spoliana 2, Aylmer, Que., distinct pattern; (12) mali 6 paratype, Bell’s Corners, Ont., 
diffuse pattern; (13) mali 2? paratype, Bell’s Corners, Ont., distinct pattern; (14) oregonana 
6, S. Ottawa, Ont., diffuse pattern; (15) oregonana 2, Aweme, Man., distinct pattern; (16) 
kalmiana 6 paratype, Halifax, N.S.; (17) maracana 6, Shiawassee Co., Mich.; (18) vaccinii 
holotype 2; (19) habrosana 6, San Luis Obispo Co., Calif., diffuse pattern; (20) habrosana 
?, O'Brien, Calif., distinct pattern; (21) senatrix 2 paratype, Cochise Co., Ariz.; (22) 
costomaculana 9°, Dryden, N.Y.; (23) virginiana 4, Ithaca, N.Y. 


VOLUME 40, NUMBER 3 229 





Fics. 24-40. Pseudexentera male genitalia, some images reversed. (24) cressoniana, 
Livingston Co., Mich., prep. MAM 314792; (25) faracana, Mt. Airy, Pa., prep. WEM 
137844; (26) sepia, Kansas City, Mo., prep. SMG 905822; (27) hodsoni, Bellaire, Tex., 
prep. ECK 817; (28) knudsoni, Comal Co., Tex., prep. ECK 325; (29) haracana, Lake- 
hurst, N.J., prep. WEM 197848; (30) oreios, Sta. Cruz Co., Ariz., prep. WEM 307851; 
(31) spoliana, Blain, Pa., prep. VA 265; (32) mali, Oneida Co., Wis., prep. VA 106; (33) 
oregonana, Meach Lk., Que., prep. WEM 97851; (34) kalmiana, Mer Bleue, Ont., prep. 
WEM 135853; (35) maracana, Livingston Co., Mich., prep. JAB 80; (36) vaccinii, Ingham 
Co., Mich., prep. JAB 79; (37) habrosana, Alameda Co., Calif., prep. WEM 257852; (38) 
senatrix paratype, Cochise Co., Ariz., prep. WEM 237844; (39) costomaculana, Ithaca, 
N.Y., prep. WEM 251852; (40) virginiana, Ithaca, N.Y., prep. WEM 251854. 


Diagnosis. Forewing pattern varying within sexes, overlapping between sexes, females 
averaging higher in contrast (Figs. 14, 15) (157n). Forewing veins R, and R; usually 
(92%) stalked or connate at origin, sometimes (8%) approximate (37n). In males, valva 
constricted approximately at middle, valval length/cucullus length ratio 1.9 to 2.2, anal 
spine near lower edge of cucullus, lower edge of cucullus lacks projections, aedeagus has 
falcate apex (Fig. 33) (14n). In females, ostium bursae begins % to 1% its width behind 
front edge of sternum, forward end of sterigma tapers gradually if at all, corpus bursae 
spicule bases nowhere fused into a sclerotized patch, signa unequal or subequal in size 
(Fig. 50) (18n). Forewing length of males 7.5 to 10.0 mm (85n), of females 7.5 to 9.5 
mm (58n). 

Comments. I examined adults from Alberta, British Columbia, Maine, Manitoba, Mich- 
igan, Nova Scotia, Ontario, Oregon, Quebec, Saskatchewan, and Wisconsin (BMNH, 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 41-57. Pseudexentera female genitalia. (41) cressoniana, Barry Co., Mo., prep. 
SMG 1109824; (42) faracana, Putnam Co., Ill., prep. DH 617812; (48) sepia, Putnam 
Co., Ill., prep. DH 630812; (44) hodsoni, Oktibbeha Co., Miss., prep. SMG 1029821; (45) 
knudsoni, San Jacinto Co., Tex., prep. WEM 35852; (46) haracana, Vicksburg, Miss., 
prep. SMG 923821; (47) oreios, Cochise Co., Ariz., prep. WEM 35853; (48) spoliana, 
Columbia, Mo., prep. SMG 1206821; (49) mali, Ithaca, N.Y., prep. WEM 304852; (50) 
oregonana, Ottawa E., Ont., prep. WEM 95854; (51) kalmiana, Constance Bay, Ont., 
prep. WEM 218851; (52) maracana, Putnam Co., Ill., prep. DH 701814; (58) vaccinii, 
Washtenaw Co., Mich., prep. VA 252; (54) habrosana, O’Brien, Calif., prep. WEM 
105853; (55) senatrix, Yavapai Co., Ariz., prep. WEM 119851; (56) costomaculana, New 
Brighton, Pa., prep. WEM 301858; (57) virginiana, Pittsburgh, Pa., prep. WEM 116852. 


VOLUME 40, NUMBER 3 Dee | 


CNC, LACM, MSUE, NSM, UCB, UMMZ). The study sample included the lectotype, a 
paralectotype, adults whose wings were illustrated by Freeman (1942) (3n), and adults 
reared from Populus tremuloides Michx. (28n) and Salix sp. (15n). 

Biology. The larva eats new foliage and rolls leaves of Populus tremuloides and Salix 
sp. There is one generation per year, with the pupa wintering on the ground (Freeman 
1942, MacKay 1962, McDunnough 1940, Prentice 1966, Wong & Melvin 1967). Adult 
capture dates range from 15 March to 14 June (96n). 


Pseudexentera kalmiana McDunnough 
(Figs. 16, 34, 51) 


Pseudexentera kalmiana McDunnough (1959:4) (holotype male: White Point Beach, 
Queens Co., Nova Scotia, reared from Kalmia sp. [angustifolia L. according to 
Ferguson 1975] 15 April 1954, J. McDunnough, genit. prep. WEM 148851, forewing 
length 6.0 mm, No. 6807 in CNC). 


Diagnosis. Forewing pattern (Fig. 16) varying little between or within sexes (58n). 
Forewing veins R, and R, usually (80%) connate or approximate at origin, sometimes 
(20%) stalked or separate (34n). In males, valva constricted at middle, valval length/ 
cucullus length ratio 1.8 to 1.9, anal spine near lower edge of cucullus, lower edge of 
cucullus lacks projections, aedeagus has falcate apex (Fig. 34) (14n). In females, ostium 
bursae begins % to 1% its width behind front edge of sternum, forward end of sterigma 
tapers gradually if at all, corpus bursae spicule bases nowhere fused into a sclerotized 
patch, signa subequal in size (Fig. 51) (16n). Forewing length of males 5.0 to 6.5 mm 
(36n), of females 5.5 to 6.5 mm (22n). 

Comments. | examined adults from Michigan, Newfoundland, New York, Nova Scotia, 
and Ontario (CNC, CU, MSUE, NMS). The study sample included paratypes (4n) and 
holotype. 

Biology. The larva feeds on Kalmia angustifolia L. (Ferguson 1975). Adult capture 
dates range from 11 April to 14 June (57n). 


Pseudexentera maracana (Kearfott) 
(Figs. 17, 35, 52) 


Proteopteryx maracana Kearfott (1907:46) (lectotype male: Cincinnati, Ohio, 3 April 
1906, A. F. Baun, selected by C. Heinrich, designated by Klots 1942, in AMNH). 

Exentera maracana; Heinrich (1923:177). 

Pseudexentera maracana; Powell (1983:36). 


Diagnosis. Forewing pattern (Fig. 17) varying little between or within sexes, females 
possibly averaging higher in contrast (24n). Forewing veins R, and R, usually (95%) 
stalked or connate at origin, sometimes (5%) approximate (22n). In males, valva con- 
stricted approximately at middle, valval length/cucullus length ratio 1.6 to 1.8, anal 
spine near lower edge of cucullus, lower edge of cucullus lacks projections, aedeagus has 
falcate apex (Fig. 85) (Qn). In females, ostium bursae begins % to 1% its width behind 
front edge of sternum, forward end of sterigma tapers gradually if at all, corpus bursae 
spicule bases nowhere fused into a sclerotized patch, signa unequal or subequal in size 
(Fig. 52) (11n). Forewing length of males 7.0 to 7.5 mm (12n), of females 6.0 to 7.0 mm 
(12n). 

Comments. I examined specimens from Illinois, Michigan, Minnesota, Mississippi, Mis- 
souri, New York, Ohio, Pennsylvania, Quebec, and Texas (AMNH, CNC, Craig, CU, 
Heitzman, INHS, Knudson, LACM, Mather, MSUE, UCB, UMMZ). The study sample 
included the lectotype, a paralectotype, and adults reared from Crataegus sp. (4n). 

Biology. The larva feeds on Crataegus. Adult capture dates range from 30 January to 
15 May (20n). 


932 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pseudexentera vaccinii Miller, new species 


(Figs. 18, 36, 53) 


Diagnosis and description. Forewing pattern (Fig. 18) varying little between or within 
sexes, females possibly averaging higher in contrast (86n). Forewing veins R, and R, 
usually (60%) connate at origin, sometimes (40%) approximate or stalked (30n). In males, 
valva constricted approximately at middle, valval length/cucullus length ratio 1.8 to 2.0, 
anal spine near lower edge of cucullus, lower edge of cucullus lacks projections, aedeagus 
has faleate apex (Fig. 36) (19n). In females, ostium bursae begins % to 1% its width 
behind front edge of sternum, forward end of sterigma tapers gradually if at all, corpus 
bursae spicule bases nowhere fused into a sclerotized patch, signa subequal in size (Fig. 
53) (16n). Forewing length of males 6.5 to 8.0 mm (21n), of females 6.0 to 7.5 mm (17n) 
(holotype 7.0 mm). Head. Labial palpus brownish white, paler on inner side, second 
segment subequal in length to eye diameter, apical segment % length of second; front 
and crown brownish white. Thorax. Hues similar to front and crown dorsally, shiny 
white ventrally; front and middle legs mixed white and brown, hind legs paler, tarsi of 
all legs banded; forewing upper-side dark markings near tawny (38) and fuscous (21), 
underside pale brown; hindwing upper side pale brown, underside paler. Abdomen. 
Brownish white dorsally, paler ventrally. Male Genitalia. Vesica with 17 to 23 deciduous 
cornuti (11n). 

Type data. Holotype female (Fig. 18): S. March, ONTARIO, 22 April 1944, J. Mc- 
Dunnough, reared from Vaccinium sp., genit. prep. Exen 1la (CNC). Thirty-seven para- 
types (ANSP, CNC, LACM, MSUE, NSM, UMMZ, UMSP): MASSACHUSETTS: Barnsta- 
ble, 22 June 1951, C. P. Kimball, 2 genit. prep. Pseud. B; same data except 8 July 1950, 
6 genit. prep. Pseud. B; MICHIGAN: Ingham Co., 8 April 1967, J. P. Donahue, ¢ genit. 
prep. JAB 76; same data except 6 genit. prep. 71; same data except 12 April 1968, 4; 
same data except 2 genit. prep. WEM 87853; same data except 2 genit. prep. JAB 69; 
same data except 6 genit. prep. VA 277; same data except 6 genit. prep. VA 278; same 
data except 6 genit. prep. VA 279; same data except 15 April 1967, 6 genit. prep. JAB 
79 (Fig. 36); same data except 26 April 1970, 2 genit. prep. HS 3177311; same data 
except 2 May 1968, 2 genit. prep. WEM 87851; same data except 18 May 1966, 2 genit. 
prep. PJ 61; Chippewa Co., 25 May 1966, J. P. Donahue, 2 genit. prep. PJ 54; same data 
except 6 genit. prep. PJ 56; Otsego Co., 3 May 1970, J. P. Donahue, ¢ genit. prep. HS 
317787; Crawford Co., 28 April 1951, R. R. Dreisbach, 6 genit. prep. RBM 198; Wash- 
tenaw Co., 14 April 1936, W. W. Newcomb, @ genit. prep. VA 252 (Fig. 53); Midland 
Co., 21-31 May 1961, R. R. Dreisbach, 2 genit. prep. PJ 234; MINNESOTA: Cass Co., 
14 May 1936, R. H. Daggy, 2 genit. prep. ACC 427781; NEW JERSEY: New Lisbon, 24 
April 1933, E. P. Darlington, 2 genit. prep. KL 191; same data except 6 May, 2 genit. 
prep. WEM 1022735; same data except 6 genit. prep. WEM 218853; same data except 
14 May, 6 genit. prep. WEM 1022734; same data except 16 May, 6 genit. prep. KL 196; 
same data except 14 May 1932, 6 genit. prep. KL 195; ONTARIO: Same data as holotype 
except 2 genit. prep. WEM 85853; Bell’s Corners, 20 April 1941, T. N. Freeman, 92; same 
data except ? genit. prep. LKM 824769; same data except 25 April, 6 genit. prep. WEM 
87855; Constance Bay, 28 April 1941, J. McDunnough, 6 genit. prep. WEM 218852; same 
data except T. N. Freeman, 6; same data except 6 genit. prep. Exen 11; same data except 
26 April 1935, W. J. Brown, 6 genit. prep. WEM 87854; same data except G. S. Walley, 
6 genit. prep. WEM 177841; PENNSYLVANIA: Allegheny Co., 10 April 1910, F. Marloff, 
2 genit. prep. WEM 97853. 

Comments. This species most resembles Pseudexentera maracana. It differs in its 
thinner middle forewing crossband (Figs. 17, 18), its statistically different R, and R, 
origins, its statistically different ostium bursae position and in larval host. The frequency 
distribution of approximate, connate, and stalked R, and R, in P. vaccinii is 7, 18, and 
5, respectively, compared to 1, 10, and 11 in P. maracana (G,,, = 10.4, Pa < 0.01). In 
P. vaccinii the ostium bursae begins on average 0.58 its width behind the front edge of 
the sternum compared with 0.95 in P. maracana. The difference, 0.37, is significant (t = 
3.50, Pa < 0.01). The unnamed Pseudexentera larva on Vaccinium described by MacKay 
(1959:121) is probably this species. 


VOLUME 40, NUMBER 3 233 


Biology. The larva feeds on Vaccinium (2n). Adult capture dates range from 8 April 
to 8 July (36n). 


Pseudexentera habrosana (Heinrich) 
(Figs. 19, 20, 37, 54) 


Exentera habrosana Heinrich (1923:178) (holotype male: San Diego, Calif., 17 March 
1912, W. S. Wright, genitalia illustrated by Heinrich 1923, No. 24833 in USNM). 
Pseudexentera habrosana; Powell (1961:203). 


Diagnosis. Forewing pattern varying within sexes, overlapping between sexes, females 
possibly averaging higher in contrast (Figs. 19, 20) (59n). Forewing veins R, and R, 
usually (97%) approximate or connate at origin, sometimes (3%) stalked (56n). In males, 
valva constricted approximately at middle, valval length/cucullus length ratio 1.8 to 2.1, 
anal spine located near lower edge of cucullus, lower edge of cucullus may have up to 
two inconspicuous projections ranging in shape from bumps to spinelets, aedeagus has 
faleate apex (Fig. 37) (20n). In females, ostium bursae begins % to 1% its width behind 
front edge of sternum, forward end of sterigma tapers gradually if at all, corpus bursae 
spicule bases nowhere fused into a sclerotized patch, signa subequal or unequal in size, 
forward end of sterigma tapers gradually if at all, corpus bursae spicule bases nowhere 
wing length of males 7.5 to 9.5 mm (35n), of females 7.0 to 8.5 mm (24n). 

Comments. [ examined adults from the California counties of Alameda, Contra Costa, 
Lake, Los Angeles, Marin, Mendocino, Orange, Placer, Riverside, San Diego, San Fran- 
cisco, San Luis Obispo, Santa Barbara, Santa Clara, Shasta, Sonoma, Stanislaus, Tuolumne, 
and Ventura (LACM, Leuschner, UCB, USNM). The study sample included adults listed 
by Powell (1961) (6n), adults reared later by Powell from Quercus agrifolia Née or Q. 
wislizeni A. DC. (12n), and a paratype. The larva was described by Powell (1961). 

Biology. The larva feeds on Q. agrifolia and Q. wislizeni foliage. Adult capture dates 
range from 21 January to 17 April (47n). 


Pseudexentera senatrix (Heinrich) 
(Figs. 21, 38, 55) 


Exentera senatrix Heinrich (1924:390) (holotype male: Paradise, Cochise Co., Ariz., 8— 
15 March, in USNM). 
Pseudexentera senatrix; Powell (1983:36). 


Diagnosis. Forewing pattern (Fig. 21) varying little within or between sexes, females 
possibly averaging higher in contrast (25n). Forewing veins R, and R, usually (90%) 
connate or stalked at origin, sometimes (10%) approximate (20n). In males, valva con- 
stricted approximately at middle, valval length/cucullus length ratio 2.0 to 2.1, anal 
spine near lower edge of cucullus, lower edge of cucullus may have up to four incon- 
spicuous projections ranging in shape from bumps to spinelets, aedeagus has falcate apex 
(Fig. 38) (6n). In females, ostium bursae begins % to 1% behind front edge of sternum, 
forward end of sterigma tapers gradually if at all, corpus bursae spicule bases nowhere 
fused into a sclerotized patch, signa unequal or subequal in size, forward and rear halves 
of papillae anales subsymmetrical in outline (Fig. 55) (7n). Forewing length of males 8.0 
to 8.5 mm (16n), of females 8.0 to 8.5 mm (9n). 

Comments. I examined adults from Arizona and San Bernardino Co. (Barnwell, New 
York Mts.), California (Leuschner, LACM, UCB, USNM). The study sample included 
paratypes (3n). 

This species has not been adequately differentiated from any congener. It most resem- 
bles P. habrosana. It differs statistically in R, and R, origin, in size as well as symmetry 
of papillae anales, and in other ways. The frequency distribution of approximate, con- 
nate, and stalked R, and R, in P. senatrix is 2, 10, and 8, respectively, compared to 33, 
21, and 2 in P. habrosana (G,,, = 22.0, Pa < 0.005). Length x width maxima of one 
papilla analis ranges from 0.14 to 0.16 mm? in P. senatrix (7n) compared to 0.08 to 0.12 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mm? in P. habrosana (12n). The respective means are 0.15 and 0.10, and the difference, 
0.05, is significant with or without dividing individual values by forewing length to adjust 
for body size (t = 8.4, Pa < 0.001). Also, forewing pattern varies less in P. senatrix than 
in P. habrosana. The former has coarse-grained ashy gray forewings, with brown scaling 
in the terminal area obscure and visible only under magnification. The latter has fine- 
grained forewing coloration with brown scaling conspicuous in the dark areas and often 
elsewhere as well. The two species are allopatric, P. senatrix occurring in Arizona and 
southeastern California, and P. habrosana in many areas of California, but not in the 
southeastern part. The known larval hosts of P. habrosana do not occur in the range of 
P. senatrix, which indicates the latter has a different host or hosts. 

Biology. The larval host is unknown. Adult capture dates range from 18 March to 6 
May (25n). 


Pseudexentera costomaculana (Clemens) 
(Figs. 22, 39, 56) 


Anchylopera costomaculana Clemens (1860:349) (lectotype female: North America, no 
date, No. 139, designated by Darlington 1947, forewing length 7.0 mm, wings illus- 
trated by Miller 1973a, in ANSP). 

Batodes bipustulana Walker (1863:316) (lectotype female: “N. Amer. ..., Type...,” 
no date, genit. prep. No. 11635, selected by N. S. Obraztsov, designated here, fore- 
wing length 7.5 mm, in BMNH). 

Exentera costomaculana; Heinrich (1923:178). 

Pseudexentera costomaculana; McDunnough (1954:2), MacKay (1962:640). 


Diagnosis. Forewing pattern (Fig. 22) varying little between or within sexes, pale areas 
near yellow ocher (123C) (60n). Forewing veins R, and R, approximate or separate at 
origin (27n). In males, valva constricted at % distance between base and apex, valval 
length/cucullus length ratio 2.8 to 2.9, anal spine near middle of cucullus, aedeagus has 
unmodified apex (Fig. 39) (8n). In females, ostium bursae begins % to % its width behind 
front edge of sternum, forward end of sterigma tapers sharply, corpus bursae spicule 
bases fused on one side near ductus bursae into a lightly sclerotized patch, signa unequal 
in size (Fig. 56) (8n). Forewing length of males 6.5 to 8.5 mm (19n), of females 7.0 to 
8.0 mm (138n). 

Comments. I examined adults from Connecticut, Maryland, Michigan, Mississippi, 
New Hampshire, New York, North Carolina, Nova Scotia, Pennsylvania, and West Vir- 
ginia (LACM, MSUE, NSM, UCB, UMMZ, USNM). The study sample included adults 
reared from Hamamelis sp. (prob. virginiana L.) (8n), and lectotypes of P. costomacu- 
lana and P. bipustulana. The larva was described by MacKay (1962). 

Biology. The larva mines buds and folds leaves of Hamamelis virginiana L., and the 
pupa winters (McDunnough 1954, MacKay 1962, Ferguson 1975). Adult capture dates 
range from 9 April to 18 July (55n). 


Pseudexentera virginiana (Clemens) 
(Figs. 23, 40, 57) 


Anchylopera virginiana Clemens (1865:512) (type unknown). 
Exentera virginiana; Heinrich (1923:179). 
Pseudexentera virginiana; Miller (1973a:224). 


Diagnosis. Forewing pattern (Fig. 23) varying little between or within sexes, pale areas 
near drab (27) (46n). Forewing veins R, and R, approximate or separate at origin (22n). 
In males, valva constricted at % distance between base and apex, valval length/cucullus 
length ratio 2.5 to 2.6, anal spine near middle of cucullus, aedeagus has unmodified apex 
(Fig. 40) (6n). In females, ostium bursae begins %4 to % its width behind front edge of 
sternum, forward end of sterigma tapers sharply, corpus bursae spicule bases fused on 


VOLUME 40, NUMBER 3 235 


one side near ductus bursae into a lightly sclerotized patch, signa unequal in size (Fig. 
57) (8n). Forewing length of males 7.5 to 9.0 mm (9n), of females 7.5 to 9.5 mm (13n). 
Comments. I examined adults from Connecticut, Maryland, Michigan, Mississippi, 
New York, Pennsylvania, and South Carolina (LACM, Mather, MSUE, UCB, UMMZ, 
USNM). 
Biology. The larval host is unknown. Adult capture dates range from 7 March to 11 
May (46n). 


CONCLUSION 


Only three adults, less than 0.38% of the number studied, were not 
satisfactorily resolved to any of the above species (UCB, CNC, MSUE). 
All three are different and may represent extreme variants of named 
species or single examples of unnamed species. 

Of the 18 species previously recognized (Powell 1983), the names of 
eight appear valid, and the names of five are revised here. With the 
five new species described here, the number of Pseudexentera species 
now totals 17. Thirteen occur only east of the Great Plains, three occur 
only westward, and one is transcontinental. The following table of 
equivalents summarizes Pseudexentera species names here relative to 
those in Hienrich’s (1923) revision: 


Here Heinrich 
cressoniana (Clem. ) improbana (part) 
faracana (Kft.) faracana and spoliana (part) 
sepia n. sp. spoliana (part) 
hedsoni n. sp. spoliana (part) 
knudsoni n. sp. _ 
haracana (Kft.) haracana 
oreios Ni. Sp. = 
spoliana (Clem.) improbana (part) 
mali Freeman — 
oregonana (Wlsm.) improbana oregonana 
kalmiana McD. — 
maracana (Kft.) maracana 
vaccinii n. sp. — 
habrosana (Heinr.) habrosana 
senatrix (Heinr.) — 
costomaculana (Clem. ) costomaculana 
virginiana (Clem.) virginiana 


So far as known, larval host associations by plant family and number 
of Pseudexentera species are: Fagaceae (4), Rosaceae (2), Ericaceae 
(2), Juglandaceae (1), Salicaceae (1), and Hamamelidaceae (1). Larval 
hosts are unknown for six species. Fragmentary larval host information 
may be the single greatest deficiency hindering progress and strength- 
ening of Pseudexentera taxonomy. 


ACKNOWLEDGMENTS 


I thank curators of the sources named for specimen loans and other assistance. I am 
indebted to J. A. Powell, R. L. Brown, D. C. Ferguson, R. W. Hodges, and J. F. G. 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Clarke, who read the manuscript and made many useful suggestions, and to M. Elling 
for painstakingly typing the manuscript. D. C. Ferguson kindly served as special editor 
for this paper. 


LITERATURE CITED 


Brown, R. L. 1982. Notes on Gretchena: A new species and the synonymy of Gwen- 

_ dolina (Lepidoptera: Tortricidae). Proc. Entomol. Soc. Wash. 84:594-602. 

Busck, A. 1914. Life history of Eucosma haracana Kearfott. Proc. Entomol. Soc. Wash. 
16:150. 

CHAPMAN, P. J. & S. E. LIENK. 1971. Tortricid fauna of apple in New York (Lepidop- 
tera: Tortricidae). New York Agr. Exp. Stn. Geneva Spec. Publ. 122 pp. 

CLARKE, J. F. G. 1941. The preparation of slides of the genitalia of Lepidoptera. Bull. 
Brooklyn Entomol. Soc. 36:149-161. 

CLEMENS, B. 1860. Contributions to American lepidopterology.—No. 6. Proc. Acad. 
Nat. Sci. Phila. 1860:345-362. 

(1864) 1865. North American micro-lepidoptera. Proc. Entomol. Soc. Phila. 3: 
505-520. 

DARLINGTON, E. P. 1947. Notes on certain types of Lepidoptera described by Brack- 
enridge Clemens. Trans. Am. Entomol. Soc. 73:85-104. 

FERGUSON, D. C. 1975. Host records for Lepidoptera reared in eastern North America. 
U.S. Dep. Agr. Tech. Bull. 1521. 49 pp. 

FREEMAN, T. N. 1942. A new species of Pseudexentera from apple, with notes on allied 
species (Olethreutidae, Lepidoptera). Can. Entomol. 74:212-215. 

GROTE, A. R. 1877. A new genus of Tortricidae. Can. Entomol. 9:227. 

HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucos- 
minae of the family Olethreutidae. U.S. Nat. Mus. Bull. 123. 298 pp. 

1924. North American Eucosminae, notes and new species (Lepidoptera). J. 

Wash. Acad. Sci. 14:385-398. 

1940. Correction of a misused generic name (Lepidoptera, Olethreutidae). Can. 
Entomol. 72:242-248. 

KEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc. 
33:1-98. 

Kiots, A. B. 1942. Type material of North American microlepidoptera other than 
Aegeriidae in the American Museum of Natural History. Bull. Am. Mus. Nat. Hist. 
79:391-424. 

LITTLE, E. L. 1979. Checklist of United States trees. U.S. Dep. Agr. Handbook 541. 
375 pp. 

MacKay, M. R. 1959. Larvae of the North American Olethreutidae (Lepidoptera). 
Can. Entomol. Suppl. 10. 338 pp. 

1962. Additional larvae of the North American Olethreutinae (1) (Lepidoptera: 

Tortricidae). Can. Entomol. 94:626-643. 

1965. The larva of Epinotia criddleana Kft. and a note on Pseudexentera 
oregonana Wlshm. (Lepidoptera: Tortricidae). Can. Entomol. 97:666—668. 

McCDUNNOUGH, J. 1940. A new Pseudexentera from hickory (Eucosmidae, Lepid.). 
Can. Entomol. 72:243-244. 

1954. New microlepidoptera from the region of Halifax, Nova Scotia, with 

notes on other species. Am. Mus. Novit. 1686. 15 pp. 

1959. On some changes in nomenclature of microlepidoptera, with description 
of a new species. Am. Mus. Novit. 1954. 9 pp. 

MILLER, W. E. 1973a. Clemens types of Olethreutinae (Lepidoptera, Tortricidae). 
Trans. Am. Entomol. Soc. 99:205-234. 

1973b. Two previously unrecognized scientific names for the strawberry leaf- 

roller. Ann. Entomol. Soc. Am. 66:553-554. 

1974. Identities of taxonomically confused moths of the Eucosma agricolana 

group and description of a new species (Lepidoptera, Tortricidae). Ann. Entomol. 

Soc. Am. 67:601-604. 





























VOLUME 40, NUMBER 3 Dt 





1977. Wing measure as a size index in Lepidoptera: The family Olethreutidae. 
Ann. Entomol. Soc. Am. 70:253-256. 

1979. The genus Olethreutes: Identity corrections and description of a new 
species (Lepidoptera: Tortricidae: Olethreutinae). Ann. Entomol. Soc. Am. 72:232- 
236. 








1985. Nearctic Olethreutes: Five new synonymies, two revised statuses, and 
notes (Lepidoptera: Tortricidae). Proc. Entomol. Soc. Wash. 87:408-417. 

POWELL, J. A. 1961. Taxonomic and biological observations on Pseudexentera habro- 
sana (Heinrich) (Lepidoptera: Tortricidae). Pan-Pac. Entomol. 37:203-209. 

1983. Tortricidae, pp. 31-41. In R. W. Hodges (ed.), Check list of the Lepi- 
doptera of America north of Mexico. E. W. Classey Ltd. and Wedge Entomological 
Research Foundation, London. 284 pp. 

PRENTICE, R. M. (ED.). 1966. Forest Lepidoptera of Canada recorded by the Forest 
Insect Survey. Vol. 4. Can. Dep. For. Publ. 1142:543-840. 

SMITHE, F. B. 1975. Naturalist’s color guide. American Museum of Natural History, 
New York. 182 color swatches. 

WALKER, F. 1863. List of the specimens of lepidopterous insects in the collection of 
the British Museum. Part 28. Tortricites and Tineites. [Pp. 283-561.] 

WALSINGHAM, T. DE GREY, SIXTH EARL. 1879. Illustrations of typical specimens of 
Lepidoptera Heterocera in the collection of the British Museum. 4. North American 
Tortricidae. British Museum, London. 84 pp. 

Wonc, H. R. & J. C. E. MELVIN. 1967. The leaf roller Pseudexentera oregonana 
Wlshm. Can. Dep. For. Rural Dev. Bi-Monthly Res. Notes 23(1):3-4. 





Received for publication 14 March 1986; accepted 25 June 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 238-239 


GENERAL NOTE 


AN INTER-SUBFAMILIAL MATING INVOLVING AN ENDANGERED 
BUTTERFLY (LYCAENIDAE: LYCAENINAE AND RIODININAE) 


I report here an inter-subfamilial mating between a female Euphilotes battoides allyni 
(Shields) (Lycaeninae) and a male Apodemia mormo virgulti (Behr) (Riodininae). The 
former is commonly known as the El Segundo blue butterfly, and is recognized as en- 
dangered by the U.S. Fish & Wildlife Service. 

The mating was observed on 18 July 1984 at a remnant of the El Segundo sand dune 
system at the western end of the Los Angeles International Airport (LAX), Los Angeles 
Co., CA. During July and August 1984, I conducted a capture-recapture study of the El 
Segundo blue at LAX to collect baseline information on the butterfly’s ecological require- 
ments and to estimate its population size. My study was sponsored by the California 
Department of Fish & Game. 

The mating pair (Fig. 1) was first noted at 1232 h PDT perched about 4 cm above 
the ground on foliage of Erysimum suffrutescens (Abrams) G. Rossb. (Brassicaceae). The 
wing condition of both specimens was fresh, suggesting they were no more than a few 
days old. The crucifer was surrounded by a patch of about 15 Eriogonum parvifolium 





an "ie 


Fic. 1. The inter-subfamilial mating pair, Euphilotes battoides allyni (top) and Apo- 
demia mormo virgulti (bottom). 


VOLUME 40, NUMBER 3 239 


Sm. in Rees. (Polygonaceae), larval and primary adult foodplant of both lycaenids at 
LAX. These plants were located approximately 75 m E of the airport’s main radar 
installation in the SE portion of the dunes, where they grew just below the ridgetop of 
a sand hill at about 50 m elevation. Earlier, at 1207 h PDT, I recorded a temperature 
of 31°C, with bright, sunny skies, and a slight breeze from the W measuring about 8 
km/h. 

While I tried to photograph the mating pair, it made three short flights, moving no 
more than about 3 m with each effort. The female allyni was the active flyer. The pair 
first landed on the sandy substrate, then on a dead specimen of Erysimum suffrutescens, 
and finally on a twig of an unidentified dead plant. Copulation ceased at 1337 h PDT, 
and the female allyni immediately flew away, while the male virgulti remained mo- 
tionless for another 4 min before it resumed normal activity. 

The female allyni was marked while the pair was mating, and it was observed later 
that day nectaring and ovipositing on E. parvifolium in the same area. I collected a 
single egg and the female to obtain more eggs. During the next 3 days, the female allyni 
laid 22 eggs in confinement before expiring. Upon dissecting the female’s abdomen, I 
found a properly formed spermatophore. The eggs were examined microscopically and 
compared to fertile allyni eggs. All eggs were typical of allyni; however, none were 
viable. I previously reared allyni and found that larvae hatch within 4 to 8 days after 
oviposition (Arnold, 1983, Univ. Calif. Publ. in Entomol. 99:1-161). I dissected all 23 
eggs about 3 weeks after they were laid, and found no evidence of developing larvae. 
Probably a breakdown in a premating, reproductive-isolating mechanism (perhaps be- 
havioral stimuli) led to this unusual mating, but apparently no hybrid offspring were 
produced due to the action of one or more postmating, reproductive-isolating mecha- 
nisms such as gametic mortality, zygotic mortality, or hybrid inviability. 


RICHARD A. ARNOLD, 50 Cleaveland Road #8, Pleasant Hill, California 94528. 


Received for publication 30 January 1986; accepted 22 April 1986. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 240-241 


BOOK REVIEWS 


BRITISH PYRALID Motus. A GUIDE TO THEIR IDENTIFICATION, by Barry Goater, 175 
pp., col. frontispiece, 8 col. pls., 12 text figs. Harley Books, Great Horklesley, Colchester, 
Essex CO6 4AH, England. Distributed by Apollo Books, Lundbyvej 36, DK-5700 Svend- 
borg, Denmark. 


This is an admirable work, a happy combination of excellence and meticulousness by 
author, illustrators, and publisher who clearly kept the user uppermost in mind. It meets 
its primary stated purpose, identification, by illustrations and descriptions. It deals with 
208 species but there are 264 individual color photographs of the moths, by Geoffrey 
Senior, supplemented by 58 drawings by Robert Dyke of genitalic and other character- 
istics of closely related species, and by structural diagrams. The descriptions of the 
distinguishing features of the species are concise. There are no keys, which anyway are 
too often inadequate excuses for failures to provide adequate figures, and they are neither 
needed nor missed. 

This book is more than merely a competent and overdue aid to identifications. More 
than most works of its kind it gives information on the distributions (including Ireland 
and the Channel Islands), habits, behavior, life histories, foodplants (with a separate index 
to these), and collecting histories, reflecting the wide natural history interests of its author. 
Such information is of value in indicating what is not known and thus needs to be found 
out. The early stages of such common species as Crambus pascuella and Scoparia am- 
bigualis are still undescribed. Species such as Scoparia ancipitella and Crambus ericella 
which have northerly distributions in Great Britain should be found in Ireland. Are the 
three alpine species that were recorded from the mountains of Scotland or northern 
England in Victorian times and not since then natives that await rediscovery? Why does 
Ostrinia nubilalis feed exclusively on Artemisia in Britain? 

In the 35 years since the publication of the previous book on these moths, the number 
of known British species went from 174 to 208. Eighteen, perhaps 20, of the 34 additions 
were introduced species, 1i of them on aquatic plants, and most of the remainder on 
stored food products. About 10 were previously unrecorded migrants or vagrants from 
continental Europe. Only two, and perhaps a third, were apparently previously overlooked 
natives. These additions are in part offset by the dozen or so species that were recorded 
in the 19th century but not since. Some of these may yet be rediscovered: the only British 
records for Acigonia cicatricella are 100 years apart. 

Changes in the abundance of individual species during the past 35 years are notewor- 
thy. Increases have been of a few established immigrants and introduced species. Some 
decreases clearly were from human activities: improvements in farming hygiene that 
have been detrimental to the barn-inhabiting or hay-feeding Aglossa spp. and Pyralis 
lienigialis, in bee-keeping to Galleria mellonella, and perhaps in hedge cutting to Nu- 
monia advenella, which likes old, uncut hawthorns. The decline in the extent of chalk 
downlands, and the damage by visitors and developers to sandy coastal areas may be 
primary reasons for decreases in at least five species of those habitats. 

Whether or not weather changes have had significant effects is not clear. It may not 
be coincidence, however, that the 30 or so species that inhabit relatively dry situations, 
as on sandhills or sandy or chalky soils or limestone pavement, appear as a whole to be 
more local and less common where they occur than the inhabitants of wet situations; or 
that the two migrants that became established within the last 35 years, Phlyctaenia 
perlucidalis and Ancylosis oblitella, like damp situations, whereas the one species that 
has not been found since 1960, Eurhodope cirrigerella (incidentally, a generic name not 
in the index) is believed to require hot, dry summers. In any event, the fact that so many 
species are capable of flying across the English Channel, and much greater distances to 
Ireland, is one indication of how quickly a long-term weather change could be followed 
by changes in the lepidopterous fauna of Britain. 


VOLUME 40, NUMBER 3 241 


The publisher’s advertisement on the book-jacket states that Mr. Goater had to be 
persuaded to write this book. Whoever did the persuading deserves our thanks and 
congratulations. 


BRYAN P. BEIRNE, Simon Fraser University, Burnaby, British Columbia V5A 1S6, 
Canada. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 241 


NORDEUROPAS PYRALIDER, DANMARKS DyRELIV. Bind 3, by Eivind Palm. Fauna Bgger, 
Kgbenhavn, Denmark, 287 pp., including 8 color plates, 264 black and white figures, 
and distribution maps of 219 species, 1986. Distributed (as well as volumes 1 and 2 of 
the series) by Apollo Books, Lundbyvej 36, DK-5700 Svendborg, Denmark. Price: Danish 
Kroner 400.00 + postage (15% discount for subscribers to the series). 


This useful Danish-language book invites comparison with the almost simultaneously 
published British Pyralid Moths, by Barry Goater. Palm’s book deals with a larger 
fauna—219 species actually recorded from Fennoscandia and immediately adjacent parts 
of the Netherlands, North Germany and Poland, with brief notes on some 50 more species 
that might be expected, compared with 209 species, including accidentals and greenhouse 
pests, on the British list. However, several species on the British list are not represented 
in the present treatment. On the whole the treatment of species is more extensive in this 
book than in Goater’s; the distributional information is more detailed and is supplemented 
by dot maps of distribution in the region and more detailed maps of the distribution in 
Denmark of species that occur there. For anglophone readers there is a “Summary” 
paragraph for each Danish species, in which geographical distribution in Denmark, 
habitat, and times and months of flight are briefly stated. Though this courtesy will be 
appreciated, one wonders why a similar summary was not given for the non-Danish 
species as well. The numerous black and white figures of genital and other structures 
and of wing patterns will be useful in distinguishing close species. The color plates are 
clear, neat, and satisfactory for recognition, but have not achieved the brilliance of those 
in Goater’s book. Palm has picked up some late species synonymy that was missed by 
Goater, but otherwise the two classifications are extremely close, a fact that will be a 
blessing for European lepidopterists. Nordeuropas Pyralider is convenient in size and 
attractively produced. It has few blemishes, but there is some typographical confusion 
on line 2 of the captions for Figs. 205-210. 

Language and geography will in large part determine the readership of these two 
books, but the specialist in Pyralidae and the general student of North European moths 
ought to have both. As a reference for purchase by libraries, museums and universities, 
Nordens Pyralider can be recommended heartily. 


EUGENE MUNROE, Honorary Research Associate, Lyman Entomological Museum and 
Research Laboratory, Macdonald Campus of McGill University, Ste. Anne de Bellevue, 
Que. H9X 1C0, Canada. 


Journal of the Lepidopterists’ Society 
40(3), 1986, 242-245 


OBITUARY 


LAURENCE REMINGTON RUPERT (1902-1978) 


In 1977, we wrote to Laurence R. Rupert to request historical information on the 
Southern Silvery Blue [Glaucopsyche lygdamus (Doubleday)] locality which he had dis- 
covered near Horseheads, Chemung County, New York, 30 years before. Over the next 
several months we had a regular correspondence, in the course of which Mr. Rupert told 
us much about his life and Lepidoptera collecting experiences. Rupert died soon after, 
and since he was a charter member of the Lepidopterists’ Society and remained a mem- 
ber until his death, we prepared this brief (and certainly incomplete) biography, drawing 
heavily from his own letters written on the dates indicated at the end of quoted passages. 
Occasionally we add explanatory insertions in square brackets. 

Laurence Remington Rupert was born southeast of Buffalo, at Sardinia, Erie Co., New 
York, on 2 July 1902. His parents were Clara Remington Rupert and Asahel Rupert. He 
grew up in this area, and early became interested in Lepidoptera: 


When I was about twelve years old—in 1914—I got the idea that is not rare among 
boys of that age, that I would like to collect butterflies. I had no sponsor, and had no 
real idea how to go at the job, but my father bought me a copy of Holland’s Butterfly 
Book. I managed to acquire a rather messy collection of about 35 species of butterflies 
that were more or less common here at the time. But I lost interest when I seemed 
unable to get any more species, and when the collection I had fell prey to Dermestids. 

However, I did get quite well acquainted with those 35 or so species, and was able 
ever afterward to identify them. I could not get interested in skippers, for I was rarely 
able to identify them from Holland’s plates. . . . 

By the early 1920's I was in college in Albany. (I graduated from the then N.Y.S.C.T. 
in 1924 [now State University of New York at Albany].) I was not collecting butterflies 
nor moths, but was, as always, interested in hiking over the country (12 July 1977). 
The sand barrens west of the city [now known as the Albany Pine Bush] were pretty 
much intact, and there were vast acres solid blue with lupine [Lupinus perennis] 
blossoms. ... There were occasional plants interspersed with white or pink flowers. .. . 
I probably saw more blooms of it at once than I ever saw of any other flower (15 June 
1977). 

Several years later I went back to Albany for some graduate work and I was by that 
time collecting moths, Geometridae in particular. I became acquainted with Al Fred- 
erick at that time and went with him a few times to the area where the lupines grew. 
But he was interested in daytime collection of butterflies, and I was not collecting 
butterflies. I do have perhaps half a dozen geometrid moths in my collection that I 
took there, but nothing really special, only species that I later collected on the acid soil 
hills at Horseheads. One of those species is Catopyrrha coloraria, which tells me that 
there must be Ceanothus growing among the lupines (12 July 1977). [Ceanothus 
americanus is frequent in the Pine Bush. ] 


In 1933-1934, Rupert attended Cornell University Graduate School in Ithaca, New 
York, where he studied mathematics. During this time he met W. T. M. Forbes and John 
G. Franclemont, and in spring 1934 he and Franclemont collected together in the Sixmile 
Creek gorge near Ithaca. Later that year he journeyed to Mount Washington: 


In the summer of 1934, Al Frederick accompanied me on a collecting trip to Mt. 
Washington. I got a few nice moths there, and I believe he was quite successful with 
the butterflies he sought. ... Later our communications gradually faded out, for we 
really had little in common (12 July 1977). [Frederick’s collection is in the American 
Museum of Natural History in New York City—Rindge (1967).] 


In autumn 1934, Rupert started working at the Horseheads High School in Horseheads, 
New York, where he taught mathematics until 1950. His collecting of moths continued 
regularly, but he also discovered a butterfly there that has been rarely encountered in 
New York, Glaucopsyche lygdamus: 


VOLUME 40, NUMBER 3 243 


At Horseheads I was collecting almost exclusively at night. But now and then, pos- 
sibly on a Sunday afternoon, I would explore new areas by day, looking for possible 
new collecting spots, and it was on one of those trips [in 1947] that I stumbled into the 
area where the Glaucopsyche were flitting about. Somehow I thought they looked a 
little odd, although I wasn’t sure. So I took a few and showed them to Dr. Forbes the 
next time I went to Ithaca. He identified them, said they were a very nice thing... . 
At one time or another I picked up quite a few, for I found there were a few moth 
collectors with whom I was regularly exchanging, who wanted them. [See Dirig & 
Cryan, in prep., for details of Glaucopsyche lygdamus in New York.] Those are prob- 
ably the only butterflies that I have collected in over 60 years [except for some on a 
1950 trip]... in New Brunswick and the Gaspé, which IJ gave to the Cornell University 
Collection (12 July 1977). (See Ferguson & Rupert 1951.) 


During his years in Horseheads he also discovered Lambdina canitiara Rupert. This 
new geometrid moth was described from four males and a female taken at Horseheads 
between 1938 and 1943 (Rupert 1944). Rupert described the type locality as follows: 


They were all taken along the base of the next hill north of the Latta Brook, east of 
the southern end of Horseheads Village. I am sure that the entire area is now occupied 
by Route 17. However, it seems as if there must be other spots among those Southern 
Tier hills where the species must occur. It did not seem to be an area unique in any 
special way. The hill was well wooded, but the trees became scatter[ed] at the base, 
and there was a small swampy area near by, where there were alders [Alnus sp.] and 
cattails [Typha sp.]. I used to get Tacparia detersata there and Sthenopis argenteo- 
maculatus. I do not believe that the swamp area had any connection with the Lamb- 
dinas. All the known species of that genus have either oak [Quercus sp.] or pine [Pinus 
sp.] as the larval food, and most related genera do too. Moreover, I never found the 
moths in the swamp, but possibly 100 feet east, toward the hill (8 August 1977). 


Mr. Rupert was particularly interested in Geometridae, subfamily Ennominae, and 
published three additional papers on this group: “A Specific Revision of the Genus Me- 
tarranthis” (1948), “A Revision of the North American Species of the Genus Plagodis” 
(1949a), and “Notes on the Group of Genera Including Lozogramma Stephens and its 
Allies” (1949b). New combinations were made and several new species were described 
in these papers. 

After leaving his teaching position in 1950, Rupert returned to Sardinia, where he 
grew gladioli commercially for a number of years. He also worked for an electronics 
company at Arcade, New York, a few miles east of Sardinia. Throughout this time he 
continued to collect moths, but a heart ailment curtailed his activities somewhat in later 
years. In summer 1977 he still had half an acre of gladioli, and exhibited at the annual 
picnic and early seedling show of the Empire State Gladiolus Society in West Elmira on 
7 August. At that time we planned to meet, and he described himself as “easily identi- 
fiable. I am 75 years old, 5 ft. 4 in. tall and weigh about 110 pounds” (25 July 1977). He 
was also the organist in the United Methodist Church in Sardinia. 

When we asked him about patronyms, he answered 


Yes, Franclemont’s form ruperti of Catocala cerogama is named for me. So is the 
species Chytonix ruperti of the Noctuidae, which was described by Jack Franclemont 
from a series of over 60 specimens that I took at Versailles, N.Y. [west of Sardinia]. 
These are distributed in museums pretty well over the country. Dr. McDunnough also 
named a variety of Zale helata with the same name ruperti, and along with it he 
named a similar looking variety of Zale duplicata, franclemonti (12 July 1977). 


Mr. Rupert died on 138 November 1978. His moth collection is now in the Cornell 
University Insect Collection. Franclemont facilitated its acquisition, and told us that it 
was housed in ca. 125 Cornell drawers, of which 60 were noctuids. There were no 
butterflies in it at the time of Rupert’s death, these having been given to Cornell via 
Forbes years before. The Cornell Insect Collection has recently been moved to the new 
John H. and Anna B. Comstock Hall, within a compacterized arrangement of cases, and 
is being reorganized. Rupert’s specimens are still in units throughout the series, awaiting 


944 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


incorporation by species. Franclemont has a map of Rupert’s favorite collecting areas (J. 
G. Franclemont, pers. comm., 10 March 1986). 

In communications with us, Rupert made some notes on collecting sites, and reflected 
upon changes in habitats and the moth fauna during his lifetime: 


A great number of specimens in my collection are labeled RICHMOND GULF, 
SARDINIA, N.Y. Richmond Gulf is a deep ravine about seven miles west of the village, 
but in the Town of Sardinia. It is not more than a quarter of a mile long. Today it is 

heavily wooded, and I find it very unproductive in recent years. But in the [19]80’s 
and [19]40’s it was much more open and produced a lot of fine moths. . . . All specimens 
labeled EAST CONCORD, N.Y. were taken on a sphagnum bog easily located on a 
map. Also, those labeled EAST ARCADE, N.Y. were taken on another bog about ten 
miles northeast of here, or were taken on an unusual, dry, acid soil area adjacent to 
the bog, and I know which are which. For example, on the dry area there are acres 
of black chokeberry (Aronia melanocarpa) which supports a population of Catocala 
praeclara, so far as I know the only ... colony in the state west of the Hudson valley 
and Adirondacks. I also have a few other records from there of species that I have 
never seen elsewhere in western New York. 

One odd record that I have is for Merolonche dolli, which I took on the ceiling of 
my classroom in Horseheads. We had had a “Parents Night” on a hot evening in May, 
and I had all the windows open. When it was all over I noticed the strange moth on 
the ceiling. I scared up a stepladder and captured it. So far as I know it is the only 
record from upstate New York. I also have a specimen of Callopistria floridensis taken 
on the front wall of the YMCA Building in Binghamton, the only New York State 
record, I believe. This was almost certainly a stray, for it was taken in October when 
certain strays are common. 

There was much of interest in Atala [volume 4]. I was definitely interested in two 
moths mentioned [by Hessel (1976)]. In 1936 I found a female Sphinx luscitiosa on a 
fencepost at Horseheads. It was so battered and worn that it could not be identified 
except by genus. I put her in a sack and she deposited about a dozen eggs before 
expiring. I had to try all the known foods of the genus, but when the young larvae 
started to eat willow [Salix sp.], I knew what species I had. I reared six moths, four 
males and two females, the only ones in my collection. I have never seen the moth 
since. An interesting thing about the larvae was that part of them were green, as one 
might expect sphingid larvae to be, but the rest were vivid purple! Callosamia pro- 
methea used to be, apparently, our commonest Saturniid [at Sardinia], but I haven't 
seen a cocoon for years. However, a friend of mine, who lives some three miles 
northwest of town, told me last fall about the horrible creatures that were eating up 
his lilac [Syringa vulgaris] bush, and that he rapidly demolished all of them. His 
description sounded like promethea larvae, although they may have been [Hyalophora] 
cecropia (18 August 1977). 

My own collecting experience has shown me how collecting spots have changed over 
the years, with the old plants and trees increasing in size, and new ones crowding in. 
Sometimes as those changes occur, moths formerly common disappear entirely. Other 
species, once quite common, seem to have disappeared everywhere. I can give an 
outstanding example in your family of butterflies. When I was a kid the second com- 
monest butterfly around here was the one Holland called Chrysophanus hypo- 
phlaeas.... It was everywhere in gardens, fields and roadsides. I haven’t even seen 
one now in many years. Pieris rapae, of course, was the commonest, and still is (15 
June 1977). 

It has been of interest to me that in recent years I have seen no signs of a number 
of species that were common, or even abundant [in my youth at Sardinia]. Using the 
nomenclature in Dr. Forbes’ Lepidoptera of New York and Neighboring States, these 
would include Papilio cresphontes, the Lycaena phlaeas americana, ... the similar 
but larger species thoé, Euptoieta claudia, Brenthis selene myrina, Aglais |-album 
j-album, and Aglais milberti. On the other hand, at least two species that I looked for 
in those days and never saw became quite common in later years, Limenitis archippus 


VOLUME 40, NUMBER 3 245 


and Lethe portlandia. This latter one was probably here all the time, for I have seen 
it only on bait spread for moths in early evening (12 July 1977). 


Mr. Rupert’s sister, Florence Rupert, still lives at Sardinia, New York. 


ACKNOWLEDGMENTS 


We thank Brenda R. Krotz (Town Clerk of Sardinia) and especially John G. Francle- 
mont for helpful biographical information on Mr. Rupert. Franclemont also kindly sup- 
plied three references and reviewed a draft of this biography before publication. 


REFERENCES CITED 


Diric, R. & J. F. CRYAN. The Silvery Blue Butterfly [Glaucopsyche lygdamus (Double- 
day), Lycaenidae] in New York State. (in preparation). 

FERGUSON, D. C. & L. R. RUPERT. 1951. The results of a collecting trip to the Gaspé 
Peninsula. Lepid. News 5:53-54. 

HESSEL, S. A. 1976. A preliminary scan of rare and endangered Nearctic moths. Atala 
4:19-21. 

RINDGE, F. H. 1967. Recent additions to the Lepidoptera collection of the American 
Museum of Natural History. J. Lepid. Soc. 21:180. 

RUPERT, L. R. 1948. A specific revision of the genus Metarranthis (Lepidoptera, Geo- 
metridae, Ennominae). J. New York Entomol. Soc. 51: 133-159. 

1944. A new species of Lambdina, and notes on two species of Besma (Lepi- 

doptera, Geometridae, Ennominae). J. New York Entomol. Soc. 52:329-333. 

1949a. A revision of the North American species of the genus Plagodis (Lepi- 

doptera, Geometridae, Ennominae). J. New York Entomol. Soc. 57:19—49. 

1949b. Notes on the group of genera including Lozogramma Stephens and its 

allies (Lepidoptera, Geometridae, Ennominae). Proc. Entomol. Soc. Wash. 51:137- 

151. 











ROBERT Diric, The L. H. Bailey Hortorium Herbarium, Division of Biological Sci- 
ences, 467 Mann Library Building, Cornell University, Ithaca, New York 14853 AND 
JOHN F. CRYAN, New York State Department of Environmental Conservation, 2 World 
Trade Center, Room 3328, New York City, New York 10047. 


Date of Issue (Vol. 40, No. 3): 23 January 1987 





ap A: Ths = daageny Mee» 
iT) olit S44 aera 


; LL ral 


y F) 


EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
BoyYcE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Short manuscripts concerning new state records, current events, and notices 
should be sent to the News, June Preston, Editor, 8832 Sunset Drive, Lawrence, KS 66044 
U.S.A. Journal contributors should prepare manuscripts according to the following in- 
structions. 

Abstract: An informative abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name with author. Mea- 
surements should be given in metric units; times should be given in terms of the 24- 
hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. References 
to footnotes should be numbered consecutively, and the footnotes typed on a separate 
sheet. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In general notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-80). 

Illustrations: Only half of symmetrical objects such as adults with wings spread 
should be illustrated, unless whole illustration is crucial. Photographs and drawings should 
be mounted on stiff, white backing, arranged in the desired format, allowing (with 
particular regard to lettering) for reduction to their final width (usually 4% inches). 
Illustrations larger than 8% x 11 inches are not acceptable and should be reduced pho- 
tographically to that size or smaller. The author's name, figure numbers as cited in the 
text, and an indication of the article’s title should be printed on the back of each mounted 
plate. Figures, both line drawings and halftones (photographs), should be numbered 
consecutively in Arabic numerals. The term “plate” should not be employed. Figure 
legends must be typewritten, double-spaced, on a separate sheet (not attached to the 
illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to 
each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer's errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the 
proofs. A page charge of $60 per full Journal page is applicable to all authors. Unfunded 
authors may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


A SIMPLE INSTANT DIET FOR REARING ARCTIIDAE AND OTHER 
MOTHS. Rainer Bergomaz ¢ Michael Boor ..:ccccccccecnene 131 


EVOLUTION AND IDENTIFICATION OF THE NEW WORLD HAIR- 
STREAK BUTTERFLIES (LYCAENIDAE: EUMAEINI): ELIOT’S 
TRICHONIS SECTION AND TRICHONIS HEWITSON. Robert K. 
Robbinis bc a 138 


IDENTITY OF “AUTOGRAPHA’’ OTTOLENGUII DYAR AND 
OCCURRENCE OF AUTOGRAPHA BURAETICA (STAUDINGER) IN 
NORTH AMERICA (NOCTUIDAE: PLUSIINAE). J. D. Lafon- 
CAINE Ne 158 

THE LOCATION OF MONARCH BUTTERFLY (DANAUS PLEXIPPUS 
L.) OVERWINTERING COLONIES IN MEXICO IN RELATION TO 
TOPOGRAPHY AND CLIMATE. William H. Calvert & Lincoln 


} P. Brower (0 oe ee re 164 
A NEW EUCHLOE (PIERIDAE) FROM NORTHWESTERN MEXICO. 
Paul A. Opler 2000 188 


_ PHYSICAL CONSTRAINTS OF DEFENSE AND RESPONSE TO INVER- 

| TEBRATE PREDATORS BY PIPEVINE CATERPILLARS (BATTUS 
PHILENOR: PAPILIONIDAE). Nancy E. Stamp 2c 191 

NATURAL HISTORY OF GNOPHAELA LATIPENNIS (BOISDUVAL) 
(ARCTIIDAE: PERICOPINAE) IN NORTHERN CALIFORNIA. 
George L. Godfrey & Laurence Crabtree li ene 206 

GROWTH OF THE BUCKMOTHS HEMILEUCA LUCINA AND H. MAIA_ 
(SATURNIIDAE) ON THEIR OWN AND ON EACH OTHER'S 


HOSTPLANTS. Nancy E. Stamp & M. Deane Bowers _. 214 
THE SPECIES OF PSEUDEXENTERA (TORTRICIDAE). William E. 
Miller) 20k OD ST a 218 


GENERAL NOTE 
An inter-subfamilial mating involving an endangered butterfly (Lycaenidae: 


Lycaeninae and Riodininae). Richard A. Armd o......cccccccccccsessssesessssseesenesnee 238 
Book REVIEWS 
British pyralid moths. A guide to their idemtifiCatiOm occ cccccccceecceeneceencereeeereeneeen 240 
Nordeuropas Pyralider, Danmarks Dyreliv, Bite 3 ocecsceccccesccccsscccsssccsessscsssesnsessseneonee 241 
OBITUARY 


Laurence Remington Rupert (1902-1978) 0000 ee 242 


2 -_ 
reine = 


3 Eee 


Volume 40 1986 Number 4 


ISSN 0024-0966 


JOURNAL 


of the 


_ LEPIDoPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


; 


Bek Sy Ba oF 
18 1D Quip mio, 
ASD UNS BESSS 





11 March 1987 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


DouGLas C. FERGUSON, President GERARDO LAMaASs M., Vice President 
CLIFFORD D. FERRIS, Immediate Past President | EBBE SCHMIDT NIELSEN, Vice 
OLAF H. H. MIELKE, Vice President President 

RICHARD A. ARNOLD, Secretary ERIC H. METZLER, Treasurer 


Members at large: 


BoycE A. DRUMMOND III MIRNA M. CASAGRANDE M. DEANE BOWERS 
JOHN LANE EDWARD C. KNUDSON RICHARD L. BROWN 
ROBERT K. ROBBINS FREDERICK W. STEHR PAUL A. OPLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists’ Society, to: Eric H. Metzler, Treasurer, 
1241 Kildale Square North, Columbus, Ohio 43229-1306, U.S.A.; and address changes to: 
Julian P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 
90007-4057 U.S.A. 


To obtain: 

Back issues of the Journal and NEWS (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society’) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, CA 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Boulevard, 
Los Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, CA and addi- 
tional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Soci- 
ety, % Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 


Cover illustration: First stage larva of Natada nasoni (Grote) (Limacodidae), from 
Dyar 1899, J. New York Entomol. Soc. 7:61-67. Suggested by Marc E. Epstein. 


JOURNAL OF 


Tue LeEpPIDOPTERISTS’ SOCIETY 


Volume 40 1986 Number 4 


Journal of the Lepidopterists’ Society 
40(4), 1986, 247-254 


PRESIDENTIAL ADDRESS, 1986: 
UNEXPLORED HORIZONS—THE ROLE OF THE 
AMATEUR LEPIDOPTERIST 


CLIFFORD D. FERRIS 


Bioengineering Program, University of Wyoming, 
Laramie, Wyoming 82071 


During the past few years, members have voiced various concerns 
about our Society, its meetings, and the nature of some papers pub- 
lished in our Journal. Comments include: 1, The Society is only for 
professionals these days. 2, The papers presented at meetings are too 
technical—where are the informal field trip-slide show presentations 
of old? 3, Only specialists can read the Journal. 4, How can an amateur 
hope to contribute or gain anything from membership in the Society? 

Before I respond to these comments, let us look at who is an amateur 
and who is a professional. One definition is that a person who is paid 
for his efforts is a professional, and a person who does the same job 
without being paid is an amateur. An alternative definition might be 
that a person with formal training (meaning a college degree) in ento- 
mology or zoology is a professional (in the context of our Society), 
while a person who lacks such training is an amateur. Perhaps it would 
be better to use the term lay person rather than amateur. By either 
definition, I am a lay person, as are many other folks sitting in this 
room. I am a scientist by training, but not formally trained as an 
entomologist or lepidopterist, and with one exception in the past, not 
paid to work only with insects. 

Among the people in this room are physicians, dentists, lawyers, 
engineers, professors and teachers, scientists from various disciplines, 
housewives, individuals who simply like butterflies and moths, young 
people, students, and yes—a few paid professional entomologists. Our 
Society is composed of a very broad spectrum of disciplines and inter- 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ests, and the membership as a whole is not much different from the 
sample here today. 

In his 1984 Presidential Address at the Alberta Meeting, Lee Miller 
singled out the myriad contributions made by amateurs to the study 
of Lepidoptera. Many of the past giants in our field were not profes- 
sional entomologists. Skinner was a physician, Scudder a librarian, 
Henry Edwards an actor, Lord Rothschild a banker, William Henry 
Edwards a coal baron, Bates a naturalist and explorer, Bean an artist, 
and the list goes on. All of these individuals had one thing in common— 
they were interested in nature. The term used in those days was nat- 
uralist. Amateur or professional had no meaning. A naturalist was sim- 
ply a person who was interested in and observed nature, and most 
naturalists were self-educated. They learned by reading, observing, and 
conversing or corresponding with other naturalists. It’s a shame this 
term has nearly fallen from use, at least in North America. 

We here are all naturalists with a common interest in Lepidoptera. 
So let us call ourselves naturalists whose major interest is lepidopter- 
ology. The only differences among us are the degrees to which we 
pursue our common interest. Some of us are field people who delight 
in collecting or photographing butterflies and moths. Some of us pursue 
rearing and life-history studies. Some of us work with nomenclature 
and classification, the field of taxonomy. Others of us look at details of 
behavior, genetics, and molecular biology as they relate to Lepidop- 
tera. A few of us dabble in each of these areas. 

Now let us look for a moment at how science and the ways of doing 
science have changed in recent years. Two major discoveries, one each 
in the physical and biological sciences, have altered our approach to 
all scientific disciplines. In 1947, the year in which our Society was 
founded, the transistor was invented, and it subsequently evolved into 
the microcircuit chip and microelectronics. Microelectronics has given 
us the tools and the ability to construct analytical instruments that 
measure physical and biological processes to a degree of accuracy and 
sophistication undreamed of by 19th and early 20th century naturalists. 
In 1958, Watson and Crick proposed the double-helix model for DNA 
(deoxyribonucleic acid), and molecular biology was firmly established. 
Knowledge of DNA structure coupled with electronic instruments de- 
signed to probe and elucidate this structure led to the deciphering of 
portions of the genetic code, and to the birth of genetic engineering 
and biotechnology. Discoveries being made in molecular biology are 
changing our ways of looking at biological phenomena, and are even 
changing our thinking about what constitutes a species and how we 
approach taxonomy and nomenclature. This should explain why some 


VOLUME 40, NUMBER 4 249 


Journal articles and meeting papers appear abstract—we are caught 
up in a bioscience revolution. Today biotechnology is the frontier of 
the biological sciences. 

Biotechnology and related research require extensive laboratory fa- 
cilities and sophisticated instrumentation. Such facilities are normally 
associated with large universities and research centers. Thus only a 
limited number of specialists are equipped to undertake studies of 
Lepidoptera at the molecular level. 

Now, what about the rest of us? Do we have any open frontiers, and 
if so, where are they? The answer is yes, they are all around us! While 
mclecular biology, with all its sophisticated methods to probe the very 
nature of life, is giving us information about similarities and dissimi- 
larities among insects based upon laboratory analysis, it tells us nothing 
about how those same insects behave in their natural environments. 

At one time, many specialists thought that species of Lepidoptera 
could be separated positively based upon such morphological differ- 
ences as color, wing maculation, and genitalic structure. More and 
more, we are finding out that this is not so. In the genus Colias, for 
example, the male genitalia are essentially identical and wing paterns 
are similar; yet in the field these butterflies clearly segregate into rec- 
ognizable groups that we call species. On the other hand, individual 
colonies of Erebia callias in North America and Erebia tyndarus in 
the Old World exhibit clearly defined polymorphism in male genitalia, 
although wing maculation remains constant. 

In the western United States, there is the Speyeria atlantis complex 
which may represent a single species with multiple subspecies, or nu- 
merous closely related but separate species. To date, even the molec- 
ular biology approach has failed to decipher this complex. In the long 
run, probably careful field observations of mated pairs of these but- 
terflies along with rearing adults from ova will resolve the species- 
subspecies question. The foregoing are just a few examples of problems 
that require further study. There are hundreds of unsolved problems 
in the moths, and in both tropical and arctic fauna. 

Our frontier is the field and our mission is to study how Lepidoptera 
behave in their natural habitats. From field observations, we now know 
in Speyeria mormonia that the unsilvered “‘clio” form is really only a 
form. Mixed pairs are regularly observed in copulo. On the other hand, 
we have yet to resolve the relation of the dark-disc Speyeria atlantis 
electa to the pale unsilvered-disc Speyeria atlantis hesperis. In many 
areas these two butterflies are sympatric. They were described over 
100 years ago, and we still don’t know their true taxonomic status. Are 
they varietal forms, subspecies, or species? There are many such prob- 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lems just with the species found within the forty-eight contiguous States 
and southern Canada. Collecting in the arctic and the tropics is un- 
covering many more unanswered questions. 

For many of us, our laboratory is the out-of-doors wherever butter- 
flies and moths occur. Our experimental work is to observe how these 
insects behave in the field and to rear them so that their life stages 
become known. We need to know flight patterns, courtship behavior, 
and life histories. Maybe then we can solve some of the many unan- 
swered taxonomic questions. Frequently collectors simply collect in- 
sects and check the species off on a list. We need to pay more attention 
to how lepidopterans behave in their natural environments. 

This is an area where nearly all of us can contribute. The only 
equipment required is a good pair of field boots or shoes, notebook, 
pen, binoculars, camera, and for some a butterfly net and other col- 
lecting paraphernalia. In many cases, the camera and binoculars can 
be omitted. The final ingredients are patience and perseverance. We 
need to know and record where lepidopterans fly, how they perch, 
their courtship patterns, how they oviposit and upon what, and their 
behavior in general. If we go into rearing, we need to record what the 
eggs look like, what the larvae look like in each instar, what they eat 
and what portions of the host are consumed, what the pupae look like 
and where they are placed. Do the larvae overwinter, or is hibernation 
as ova or pupae? These are simple and basic questions, yet they remain 
unanswered for some common and widespread species. Most lepidop- 
teran rearing requires only simple equipment. Some sort of rearing 
cage is desirable, a container for the larval foodplant, soil for species 
that pupate on or in the ground, and twigs or other substrates for 
species that pupate above ground. 

One of the most famous behavioral entomologists was J. H. Fabre 
who is known for his book Social Life in the Insect World. Fabre was 
an impecunious French naturalist who observed insect life about his 
garden; he was too poor to travel, yet his work is renowned the world 
over. Of equal stature to Fabre is Theodore D. A. Cockerell, who at 
age 20 traveled from his native England to the Rocky Mountains of 
Colorado in the hope that his tuberculosis would be arrested. His hope 
was realized and he died at the age of 82, renowned as a naturalist, 
scholar, and ultimately a professor at the University of Colorado—all 
without a university degree. 

What are the benefits of the studies mentioned above? They provide 
useful and very necessary information. It is only through such field 
observations that we will utimately understand the complex relations 
among many lepidopterans. The side benefits are recreation and just 
being in the out-of-doors close to Nature and her wonders. 


VOLUME 40, NUMBER 4 O51. 


Now I would like to share with you some of the problem species 
that have intrigued me for the past several years. My major interests 
lie with the arctic and arctic-alpine fauna, and the examples which 
follow derive from these. First is the satyrid Oeneis bore. For many 
years, the taxa bore and taygete were considered to be separate species, 
and were so treated by dos Passos in his 1964 Synonymic List. About 
a decade later, a paper of his read at an annual meeting of our Society 
suggested that these two taxa are conspecific, and some subsequent lists 
have taken this approach. The basis used to separate bore and taygete 
related to whitish veining on the ventral hind wings, present in taygete 
while absent in bore, although male genitalia appear identical. As more 
arctic material became available for study, it was evident, based on 
the discovery of mixed populations, that the white veining was not a 
reliable character. The name taygete fell into synonymy under the 
older name bore. Based on my collecting in the Yukon and northern 
British Columbia in 1984 and 1985, it now appears that this situation 
is not so simply resolved. While it is true that the white-venation char- 
acter is unreliable, there are clearly two phenotypes that fly together: 
one dusky and the other brightly colored. A casual observer, seeing 
only a few specimens, might simply write off these differences as nor- 
mal variation within a geographic population. Series have been col- 
lected, however, at several locations in the Yukon and northern British 
Columbia. Flight patterns are different and no mixed-phenotype mat- 
ed pairs have been observed. Opposite sexes of opposite phenotypes 
ignore one another. Preliminary examination of the male genitalia 
indicates no difference between the two color forms. In the field, the 
butterflies behave as two separate species; in the laboratory working 
with museum specimens, we would treat them as one species after 
applying the usual taxonomic methods. 

Oeneis polixenes in the western arctic behaves in a similar manner. 
There is a dark phenotype and a pale one. I first collected two females 
of the dark form in a bog in eastern Alaska in 1971. At that time, I 
simply discounted these specimens as melanic aberrants. Several years 
ago, Jim Troubridge of Cayuga, Ontario, pointed out to me that this 
dark phenotype occurs only in odd-numbered years, while the paler 
typical polixenes is annual. His collecting led him to believe that the 
biennial dark form occurs at low elevations in forest bogs, while the 
normal form occurs in more open areas and on mountain tops. In some 
localities, both forms can be collected where there is a fairly abrupt 
transition from boggy forest to open grassy hillsides. In 1985, I found 
that the dark form is not restricted to low-elevation bogs when I col- 
lected a few examples on the summit of Pink Mountain in northern 
British Columbia. As is the case with bore, the flight patterns for the 


O52. JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


two forms are quite different and I have not yet seen mixed pairs in 
copulo. No differences in male genitalia have been detected. The dark 
polixenes is found in parts of Alaska, the Yukon Territory, and north- 
ern British Columbia. 

It is not surprising that paradoxes are turning up in the western 
Arctic. Until recently, collecting in this region was limited, and one is 
frequently inhibited by weather. New roads into previously uncollect- 
ed areas, and collectors fortunate enough to encounter hot dry weather 
at the right time of year have resulted in the discovery of some fasci- 
nating yet perplexing butterflies. On the other hand, new species and 
subspecies have turned up in heavily collected parts of America; wit- 
ness Clossiana acrocnema in 1978 along a heavily traveled hiking trail 
in southern Colorado, and C. improba harryi in 1982 in central Wy- 
oming, also along a well traveled trail. Many surprises are turning up 
in Idaho, largely through the efforts of Nelson S. Curtis of Moscow, 
Idaho. Again we see the situation of differing phenotypes and voltinism 
of a supposed single species, this time in the Coenonympha tullia 
complex. In one area of Idaho, there appears to be a univoltine pop- 
ulation on the wing between flights of bivoltine population. Only care- 
ful field observations (like those by Curtis) can elucidate this sort of 
situation. From museum material, one would simply infer an extended 
flight period of a single species. 

Elsewhere in Idaho, traditional species concepts appear to break 
down. This situation is most evident in Valley Co., where among others, 
apparent hybrids or intergrades between Speyeria atlantis and S. hy- 
daspe, Euphydryas anicia and E. chalcedona, and Colias interior and 
C. pelidne have been collected. The Colias population is particularly 
interesting in that instability of phenotype is the rule. Both wing shape 
and maculation vary widely. This population was first discovered by 
Jon Shepard of Nelson, British Columbia, and its geographic range has 
since been studied by Curtis and Ferris. It is not yet clear why Valley 
Co. fosters so many unusual butterfly phenotypes. The moths collected 
there have produced nothing unusual. 

As a final example, we return to the Arctic and the Colias hecla 
complex. Several years ago, I described Colias hecla canadensis from 
Alberta and British Columbia. A somewhat similar butterfly occurs in 
the northern Yukon and sporadically in Alaska. It is not clear if two 
species are involved, or simply temporal forms of C. hecla. In the 
Ogilvie and Richardson Mts. in the Yukon, hecla that appear early in 
the season have pale-colored males with narrow wing borders, and 
females that are on the wing at the same time are usually white or 
very pale yellow with perhaps an orange flush. As the season progresses, 
brightly marked typical hecla appear in which the females are bright 


VOLUME 40, NUMBER 4 253 


orange-yellow and the males have broad dark wing borders. Are two 
species involved, or does climate play a role? Do the early-emerging 
adults represent perhaps last instars entering hibernal diapause while 
the later-emerging adults represent penultimate-instars at the time of 
hibernal diapause? Or does the early group of pale adults represent 
larvae that passed two winters, while the later-emerging bright speci- 
mens passed only one winter as larvae? Do climate and larval devel- 
opment at the time of diapause or pupation effect selective expression 
of adult color (leucopterin rather than xanthopterin in the females)? 
At this point, I cannot answer these questions. More fieldwork is need- 
ed and probably eventual rearing of adults from ova under controlled 
conditions. 

Now we return to the questions posed at the beginning of this pre- 
sentation. In short, how does the lay person fit into the Lepidopterists’ 
Society today? I would suggest simply by being a naturalist. The fron- 
tier is still in the field as it was a century ago. We still have much to 
learn about familiar butterflies and moths, and even more to learn 
about species that occur in remote regions. Since the discovery of Clos- 
siana acrocnema a few years ago, other new species described from 
North America only include Oeneis excubitor, Erebia occulta and E. 
lafontanei, and Mitoura thornei. There are undoubtedly other un- 
known species. There is also the enigma of Erebia inuitica, known 
from a single specimen. No amount of sophisticated laboratory work 
is going to uncover uncollected new species. Only a lot of leg work in 
the field will accomplish this. Laboratory instruments will not help us 
determine the geographic distributions of the many little-known but- 
terflies and moths that occur in North America and elsewhere. In some 
tropical regions, species may become extinct without our knowing that 
they ever existed, owing to extensive destruction of virgin jungle and 
lack of collecting. 

I have described a few unsolved problems that are of interest to me. 
In some cases, analytical methods derived from molecular biology might 
answer questions, but where they have been tried (in Colias and Spey- 
eria) results so far are disappointing. : 

What we do need is more field naturalists studying behavior, life 
history, and producing distribution maps. Simply accumulating data 
but not sharing information is not enough. The contribution that a 
member of our Society can make to the group as a whole is by sharing 
discoveries. What the member then gains is the satisfaction of making 
a scientific contribution and extending our knowledge of the natural 
world. 

Some members may suggest that they are too old or have physical 
disabilities and thus cannot conduct field studies. Yet I know of some 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Society members who are in their eighties and still actively collecting 
in the field. Many years ago, I exchanged specimens with a collector 
in England, who sent me fine examples of British butterflies and moths. 
One day I received a letter from his mother informing me that my 
correspondent had died. He was in his early twenties and had been an 
invalid confined to a wheelchair, but undaunted by his physical con- 
dition he reared butterflies and moths from live material supplied by 
friends. So nearly anyone can contribute to lepidopterology. 

In closing, I give you this challenge: the natural laboratory is waiting 
for you, whether it be your garden, the mountains, the prairie, the 
desert, the arctic or the tropics. The butterflies and moths are there. 
Go find out where they live and how they live, and then share your 
findings with the rest of us. You will have the satisfaction of knowing 
that you have made a contribution to your colleagues and that you 
have extended our knowledge of Lepidoptera. I wish you Godspeed. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 255-263 


SELECTIVE OVIPOSITION BY MONARCH BUTTERFLIES 
(DANAUS PLEXIPPUS L.) IN A MIXED STAND OF 
ASCLEPIAS CURASSAVICA L. AND A. INCARNATA L. 
IN SOUTH FLORIDA 


STEPHEN B. MALCOLM AND LINCOLN P. BROWER 


Department of Zoology, University of Florida, 
Gainesville, Florida 32611 


ABSTRACT. Host plant selection by ovipositing monarch butterflies occurred in a 
mixed stand of the milkweeds Asclepias curassavica and A. incarnata in a south Florida 
pasture. Three times more immature monarchs were found on A. curassavica than on 
A. incarnata. When these numbers were balanced for biomass differences between the 
two plant species, there were 5.7 times the number of monarch immatures/dry leaf 
mass/100 m? on A. curassavica than on A. incarnata. Since A. curassavica had 36 times 
more cardenolide than A. incarnata, we suggest that the basis for selective oviposition 
by monarchs is to provide an effective cardenolide-based defense for their offspring. 


Only 27 of the 108 North American species of the milkweed genus 
Asclepias (Woodson 1954) have been recorded as larval food plants of 
monarch butterflies, Danaus plexippus L. (Table 1). This restricted use 
of host species may reflect host availability, or active host selection. If, 
as Dixon et al. (1978) suggest, ovipositing monarchs do not discriminate 
between milkweed species on the basis of their cardenolide content, 
the criteria that determine patterns of resource use are likely to be 
based on the abundance, temporal and spatial distribution, and habitat 
diversity of different Asclepias species. On the other hand, Brower 
(1961) found that ovipositing female monarchs in south central Florida 
selected A. humistrata rather than nearby A. tuberosa plants. Such 
host selection may well be influenced by variations in leaf biomass and 
morphology, qualitative and quantitative chemical defenses, and nu- 
tritive value, between Asclepias species, as Price and Willson (1976) 
have suggested for another milkweed feeding specialist. 

Recent observations near Gainesville, north Florida, in spring 1983 
and 1984, indicate that monarchs do not lay eggs on the common 
milkweeds A. tuberosa and A. verticillata and rarely lay eggs on two 
less common species, A. amplexicaulis and A. tomentosa. These four 
species have low cardenolide concentrations and are medium-sized to 
small plants (Table 2; Roeske et al. 1976). Two other common milk- 
weed species, A. humistrata and A. viridis, are heavily exploited by 
ovipositing females from early April to June (Malcolm et al. 1987). 
Interestingly, these two milkweeds contain the highest concentrations 
of cardenolides and have the largest leaf biomass of the available species 
in north Florida (Table 2). Since monarchs are well known for their 
ability to store milkweed-derived cardenolides as a defense against 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. North American Asclepias species serving as hosts of monarch butterfly 
larvae in nature. 


Asclepias 
species Location Reference 
humistrata Florida Brower (1961, 1962), Nishio (1980), Cohen and 
Brower (1982), Nishio et al. (1983), Malcolm 
et al. (1987) 
Georgia Nishio (1980), Nishio et al. (1983) 
syriaca Illinois Price and Willson (1979) 
Michigan Wilbur (1976), Malcolm and Cockrell, unpubl. 
New York Rawlins and Lederhouse (1981), Malcolm and Cock- 
rell, unpubl. 
Pennsylvania Schroeder (1976), Malcolm and Cockrell, unpubl. 
Ontario Beall (1948), Urquhart (1960), Malcolm and Cock- 
rell, unpubl. 
Wisconsin Barker and Herman (1976), Borkin (1982), Malcolm, 
Cockrell, Brower, and Brower, unpubl. 
North Dakota Malcolm and Cockrell, unpubl. 
Minnesota Malcolm and Cockrell, unpubl. 
Vermont Malcolm and Cockrell, unpubl. 
Connecticut Malcolm and Cockrell, unpubl. 
New Jersey Malcolm and Cockrell, unpubl. 
Maryland Malcolm and Cockrell, unpubl. 
Virginia Malcolm and Cockrell, unpubl. 
Missouri Malcolm and Cockrell, unpubl. 
Kansas Malcolm and Cockrell, unpubl. 
Nebraska Malcolm and Cockrell, unpubl. 
Iowa Malcolm and Cockrell, unpubl. 
Ohio Brower and Brower, unpubl. 
viridis Florida Brower, Cockrell, and Malcolm, unpubl. 
Louisiana Lynch and Martin, unpubl. 
Texas Malcolm and Cockrell, unpubl. 
Arkansas Malcolm and Cockrell, unpubl. 
Oklahoma Malcolm and Cockrell, unpubl. 
Missouri Malcolm and Cockrell, unpubl. 
Kansas Malcolm and Cockrell, unpubl. 
asperula Texas Malcolm, Cockrell, Lynch, and Martin, unpubl. 
tomentosa Florida Brower, Cockrell, and Malcolm, unpubl. 
obovata? Louisiana Malcolm and Cockrell, unpubl. 
Texas Malcolm and Cockrell, unpubl. 
curassavica Florida Brower (1961), this paper 
incarnata Florida Brower (1961), this paper 
Kansas Malcolm and Cockrell, unpubl. 
Wisconsin Brower and Brower, unpubl. 
longifolia Louisiana Riley, Lynch, and Martin, unpubl. 
hirtella Arkansas Malcolm and Cockrell, unpubl. 
Missouri Malcolm and Cockrell, unpubl. 
viridiflora Michigan Wilbur (1976) 
Louisiana Lynch and Martin, unpubl. 
Texas Malcolm and Cockrell, unpubl. 
Kansas Malcolm and Cockrell, unpubl. 
amplexicaulis Florida Brower, Cockrell, and Malcolm, unpubl. 
Illinois Price and Willson (1979) 





VOLUME 40, NUMBER 4 


Asclepias 
species Location Reference 

Texas Malcolm and Cockrell, unpubl. 

Louisiana Malcolm and Cockrell, unpubl. 

Oklahoma Malcolm and Cockrell, unpubl. 
tuberosa Florida Brower (1961, 1962) 

Illinois Price and Willson (1979) 

Michigan Wilbur (1976) 
verticillata Illinois Price and Willson (1979) 

Kansas Malcolm and Cockrell, unpubl. 

Minnesota Malcolm and Cockrell, unpubl. 
exaltata Michigan Wilbur (1976) 

Virginia Malcolm and Cockrell, unpubl. 
variegata Texas Malcolm and Cockrell, unpubl. 
purpurascens ir Urquhart (1960) 

Kansas Malcolm and Cockrell, unpubl. 
lanceolata ? Urquhart (1960) 

Florida Brower, unpubl. 
sullivantii ip Urquhart (1960) 
oenotheroides Texas Lynch and Brower, unpubl. 
fascicularis California Dixon et al. (1978) 
eriocarpa California Brower et al. (1982) 
speciosa California Brower et al. (1984b) 
californica California Brower et al. (1984a) 
erosa California Brower et al., in prep. 
cordifolia California Brower et al., in prep. 
vestita California Brower et al., in prep. 


TABLE 1. Continued. 


257 


predators (Brower 1984), cardenolides may be implicated in some form 
of host selection. 

To test host selection by D. plexippus between Asclepias species, 
based on biomass and cardenolide measures, we counted the numbers 
of monarch eggs and larvae on plants within a mixed stand of two 
Asclepias species, A. curassavica and A. incarnata, that are known to 
have different cardenolide concentrations (Roeske et al. 1976). 


METHODS 


The study site was a large mixed stand of A. curassavica and A. 
incarnata in a wet pasture adjacent to a man-made lake 15 km NW 
of Miami, Dade Co., Florida (25°45'N, 80°22'W, near junction of high- 
ways US-27 and I-95). On 1 and 2 September 1984, six 10 m x 10 m, 
randomly selected plots were searched and the following measure- 
ments made: 1) number of plants per plot, 2) plant height, 3) number 
of stems per plant, 4) presence of flower buds and flowers (no seed 
pods were found), 5) number of leaf pairs per plant (like most milk- 


258 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Leaf cardenolide concentration and plant size of five milkweed species in 
May 1983 and 1984 within 25 km of Gainesville, Florida. 


Cardenolide concentration 


.l1 g dry leaf 
ug /01 € spgleay Mean dr 


geal Sa Ee Se ee Se leaf 
Asclepias species Mean SD Range N biomass/ plant (g) 
Common 
humistrata A471 157 182-797 29 Tes 
_ viridis 478 136 316-676 W 12.5 
verticillata 14 — — 1 =< (al 
Occasional 
amplexicaulis 8 3 0-6 4 1e2 
tomentosa ILS) We, 6-23 2. 1.8 


weeds these two species have opposite leaves), and 6) numbers of mon- 
arch eggs and larvae by instar. All eggs were collected and kept until 
larval emergence to determine whether they were D. plexippus or the 
queen, D. gilippus. The three eggs on A. curassavica and one on A. 
incarnata that proved to be queens were excluded from the analysis. 
Arbitrarily selected leaf samples were also collected from five plants 
of each species to measure leaf length, width and dry weight. These 
dried leaves were ground, mixed, and 0.2 g of each species extracted 
with ethanol to estimate their cardenolide concentrations by spectroas- 
say (Brower et al. 1975, 1984b). 


RESULTS 


The cardenolide concentration of A. curassavica leaves was 864 ug 
cardenolide/0.1 g dry ieaf, and that of A. incarnata 24 ug cardenolide/ 
0.1 g dry leaf. Thus A. curassavica at this location had, on average, 36 
times more cardenolide than A. incarnata. 

The six 100 m? plots contained 182 A. curassavica plants, with 430 
stems on which 33 monarch immatures were found, and 77 A. incar- 
nata plants, with 393 stems bearing 11 immature monarchs, distributed 
between the plots as shown in Table 3. 

The two milkweed species are very similar in appearance, bearing 
similar sized, lanceolate leaves. Neither leaf length, width, or shape 
(length/width) of the two species were significantly different (Table 
4a), but the dry leaves of A. incarnata were significantly heavier than 
A. curassavica. Since A. incarnata plants were significantly taller with 
more stems per plant than A. curassavica (Table 4a), they also had 
significantly more leaves (Table 4b). Thus each A. incarnata plant had 
greater biomass available to monarch larvae than A. curassavica. How- 
ever there were more than twice as many A. curassavica than A. in- 
carnata plants per 100 m? (Table 4b), which resulted in similar num- 


VOLUME 40, NUMBER 4 259 


TABLE 3. Distribution of immature monarchs on six 100 m? plots in a mixed stand 
of A. curassavica and A. incarnata near Miami, Florida, on 1 and 2 September 1984. 


No. of insects 


Instar no. 


Plot No. No. SOS on Ay hn AU EEC I, Ri aa a tg TE a 
no. Asclepias species plants stems Eggs 1 2 8 4 5 Total 
JI curassavica 34 92 6 — oy — — — 8 
it incarnata 1S} 86 1 — — — — ] 2 
2 curassavica 62 162 6 — 1 — — — Fi 
2, incarnata 16 89 — — — — — — 0 
3 curassavica 7 44 il — 1 — — — 2 
3 incarnata 8 43 — — li — — — 1 
4 curassavica 31 69 5 — — — —_ — 5 
4 incarnata 6 36 1 — — — — — jl 
5 curassavica 7, aT — — ih — — — if 
5 incarnata 18 66 — — 1 — — — ] 
6 curassavica oll 36 4 3 2, — — 1 10 
6 incarnata 16 US) 5) — ] — — — 6 
Total curassavica 182 430 22, 3 7 = — 1 33 
Total incarnata a 393 7 = 3 — = 1 11 


bers of stems overall (Table 3, t = 0.28, P = 0.78 NS) with almost 
equal leaf density and dry leaf biomass of each Asclepias species avail- 
able to ovipositing monarchs (Table 4b). 

Despite the similarity of the leaf biomass for each milkweed species, 
three times the number of immature monarchs (eggs to fifth instars) 
were found on A. curassavica than on A. incarnata (Table 4b). Simi- 
larly, when numbers of monarch immatures are corrected for host 
biomass, there were more than five times the number of monarch 
immatures/dry leaf mass on A. curassavica than on A. incarnata (Ta- 
ble 4b). 

The difference between the numbers of monarch immatures on the 
two Asclepias species is unlikely to be explained by flower attraction 
since significantly more A. incarnata plants (with fewer monarch im- 
matures) were flowering than A. curassavica (Table 4b), although most 
plants of both species were flowering. 


DISCUSSION 


We suggest that our observation of significantly more immature 
monarchs/leaf mass on A. curassavica than on A. incarnata may be 
explained by the 86 times greater cardenolide concentration of A. cur- 
assavica over A. incarnata. These results contrast with those reported 
by Dixon et al. (1978) who suggest that monarchs oviposit on Asclepias 
species with the lowest cardenolide concentrations. They found that 


260 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Plant characteristics of A. curassavica and A. incarnata on six plots in a 
mixed stand at the study site near Miami, Florida, on 1 and 2 September 1984. Differ- 
ences were tested at the 0.05 level. 


a) Leaf measurements and plant height 


Asclepias Leaf length Leaf width Leaf Leaf dry Plant height 
species (mm) (mm) length/width weight (mg) (cm) No. stems/plant 
curassavica 
Mean 81.1 14.2 5.8 34 70.5 2.4 
SE DoS) 0.6 (0), Dy 1.1 0.1 
N 20 20 20 20 182 182 
incarnata 
Mean 91.6 15.1 6.3 DO 91.1 6 J 
SE 5.8 1.0 0.4 9 1.8 0.4 
N 18 18 18 18 C0 Wil 
t 1.6 0.9 1.0 oO 9.8 8.1 
df 24 36 26 20 257 257 
IP 0.06 0.20 0.16 0.02 0.0001 0.0001 
b) Mean plant measurements and monarch numbers per 100 m? 
No. 0. No. No. mon- 
Asclepias Percent leaves/ leaves/ Leaf dr immature archs/ 
species No. plants flowering plant plot mass (g monarchs _ leaf mass 
curassavica 
Mean 29.2 64 35.2 1,148 38.5 5.5 0.17 
SE Tod ih 2.9 334 WL 1.4 0.06 
N 6 6 6 6 6 6 6 
incarnata 
Mean 12.8 90 88.2 1,130 62.5 1.8 0.03 
SE 2.0 9) 8.3 190 10.5 0.9 0.01 
N 6 6 6 6 6 6 6 
t Dn, 2.9 6.0 0.03 1.6 Phen Pappa 
df 6 10 7 10 10 10 +) 


P 0.03 0.01 0.001 0.49 0.07 0.03 0.04 


D. plexippus laid more eggs on A. curassavica, with a cardenolide 
concentration of 56 wg/0.1 g dry leaf, than on Gomphocarpus fruti- 
cosus, with 63 wg cardenolide/0.1 g dry leaf. However, their carden- 
olide concentrations are too similar to reach any conclusion as to car- 
denolide-based oviposition preference, particularly as they are the same 
cardenolide determinations first reported as “approximate amounts 
+50%”’ (Rothschild et al. 1975), and were not determined for the same 
plants used in their oviposition experiments. Gomphocarpus fruticosus 
is also an African milkweed species which casts doubt on experimental 
relevance, since monarch butterflies will only encounter recently in- 
troduced plants of this species in Australia. Using Australian D. plex- 
ippus, Zalucki and Kitching (1982) also found that females preferred 
to lay eggs on A. curassavica rather than on A. fruticosa (=G. fruti- 
cosus); but citing different published data on cardenolide concentra- 


VOLUME 40, NUMBER 4 261 


tions (Roeske et al. 1976), they suggested the reverse, that monarchs 
laid eggs on the milkweed species with most cardenolide. 

The use of milkweed-derived cardenolides appears to be at least a 
partial defense against wild avian predators for adult monarchs over- 
wintering in Mexico (Fink & Brower 1981, Fink et al. 1983, Brower 
& Calvert 1985, Brower & Fink 1985). It is likely that monarchs in 
south Florida also benefit from cardenolide-based protection against 
bird predators, particularly as late summer bird migrants pass through 
south Florida. 

Although we recently found that monarchs regulate their carden- 
olide concentrations by increasing or reducing the cardenolide concen- 
trations from their larval host plants, milkweeds such as A. incarnata 
have insufficient cardenolide from which monarchs can concentrate 
an effective cardenolide-based defense. For example, although mon- 
archs reared on A. speciosa can almost double their cardenolide con- 
centrations relative to those of host plants, from 90 to 179 ug/0.1 g 
(Brower et al. 1984b), this concentration is less than the concentrations 
in butterflies reared on other species of cardenolide-rich milkweeds. 
Roeske et al. (1976) found that adult monarchs reared from A. curas- 
savica reflect the high cardenolide concentration of their host leaves, 
in this case of 386 wg/0.1 g dry leaf, with a concentration of 319 yng / 
0.1 g dry butterfly. In contrast, they found monarchs reared from A. 
incarnata with 0-28 wg cardenolide/0.1 g had between 28 and 127 ug 
cardenolide/0.1 g dry butterfly. The emetic response of bird predators 
increases with cardenolide concentration above a lower threshold de- 
pendent on cardenolide polarity (Roeske et al. 1976). Thus monarchs 
that fed on A. curassavica as larvae at our site near Miami will be 
much better protected by cardenolides against bird predators than 
monarchs that developed on A. incarnata. 

Since individual plants of both species have a mean dry biomass 
sufficient to support the development to pupation of at least one mon- 
arch larva (between 0.92 g and 1.74 g dry leaf is required [Schroeder 
1976, Dixon et al. 1978]; A. curassavica has 0.0384 xX 35.21 = 1.20 g, 
and A. incarnata has 0.055 x 88.23 = 4.87 g [Table 4]) a monarch 
larva need not move from plant to plant, either within, or between the 
two milkweed species. Thus the effectiveness of cardenolide-based 
monarch defense is likely to be determined primarily by the oviposi- 
tion behavior of the adult female rather than by larval movements, 
particularly in view of the unpredictable costs and benefits of such 
larval movement between milkweeds (Borkin 1982). Unlike Borkin, we 
do not find instars 2 and 3 moving between plants in Florida. If late 
instars move between plants, they are more likely to find another A. 
curassavica plant. Their feeding experience may also ensure that they 
keep moving until they find another host plant of the same species. 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cardenolides are unlikely to be the sole determinant of host selection 
in these observations. Other explanations of the observed oviposition 
bias for A. curassavica include 1) females may in some way be able to 
perceive a nutritional superiority of A. curassavica over A. incarnata 
(Erickson 1973); 2) monarch females may be more attracted to the 
orange and yellow flowers of A. curassavica than the pink flowers of 
A. incarnata (however, in central Florida monarchs preferred to ovi- 
posit on the pink flowered A. humistrata compared to the orange 
flowered A. tuberosa (Brower 1961, 1962)); and 3) females may have 
responded to the number of plants available for each species (Table 3) 
rather than to the similar quantities of stems, leaves, and leaf biomass. 

Nevertheless, whatever the explanation, the natural experiment de- 
scribed in this paper as well as the observations reported by Brower 
(1961, 1962) are evidence for the oviposition preference by monarch 
butterflies for cardenolide-rich milkweed species. We suggest that this 
choice may be the result of natural selection having favored a discrim- 
inatory mechanism allowing adult female monarchs to choose milk- 
weed species that provide their offspring with a more effective car- 
denolide-based defense against predators. 


ACKNOWLEDGMENTS 


Jim Anderson and Andrew Brower helped collect the field data, and the paper was 
improved by the critical comments of Barbara Cockrell, M. P. Zalucki and an anonymous 
reviewer. 


LITERATURE CITED 


BARKER, J. F. & W. S. HERMAN. 1976. Effect of photoperiod and temperature on 
reproduction of the monarch butterfly Danaus plexippus. J. Insect Physiol. 22:1565- 
1568. 

BEALL, G. 1948. The fat content of a butterfly Danaus plexippus Linn. as affected by 
migration. Ecology 29:80-94. 

BoRKIN, S. S. 1982. Notes on shifting distribution patterns and survival of immature 
Danaus plexippus (Lepidoptera: Danaidae) on the food plant Asclepias syriaca. 
Great Lakes Entomol. 15:199-206. 

BROWER, L. P. 1961. Studies on the migration of the monarch butterfly 1. Breeding 
populations of Danaus plexippus and D. gilippus berenice in south central Florida. 
Ecology 42:76-83. 

1962. Evidence for interspecific competition in natural populations of the 

monarch and queen butterflies Danaus plexippus and D. gilippus berenice in south 

central Florida. Ecology 48:549-552. 

1984. Chemical defense in butterflies. Symp. R. Entomol. Soc. London 11:109- 








134. 

BROWER, L. P. & W. H. CALVERT. 1985. Foraging dynamics of bird predators on 
overwintering monarch butterflies in Mexico. Evolution 39:852-868. 

BROWER, L. P., M. EDMUNDS & C. M. MorFiTT. 1975. Cardenolide content and pal- 
atability of Danaus chrysippus butterflies from West Africa. J. Entomol. (A) 49:183- 
196. 

BROWER, L. P. & L. S. FINK. 1985. A natural toxic defense system: Cardenolides in 
butterflies versus birds. Ann. N.Y. Acad. Sci. 443:171-188. 


VOLUME 40, NUMBER 4 263 


BROWER, L. P., J. N. SEIBER, C. J. NELSON, S. P. LYNCH, M. P. HOGGARD & J. A. COHEN. 
1984a. Plant-determined variation in cardenolide content and thin-layer chroma- 
tography profiles of monarch butterflies, Danaus plexippus, reared on the milkweed, 
Asclepias californica in California. J. Chem. Ecol. 11:1823-1857. 

BROWER, L. P., J. N. SEIBER, C. J. NELSON, S. P. LYNCH & M. M. HOLLAND. 1984b. 
Plant-determined variation in the cardenolide content, thin-layer chromatography 
profiles, and emetic potency of monarch butterflies, Danaus plexippus L. reared on 
milkweed plants in California: 2. Asclepias speciosa. J. Chem. Ecol. 10:601-639. 

BROWER, L. P., J. N. SEIBER, C. J. NELSON, S. P. LYNCH & P. M. TUSKES. 1982. Plant- 
determined variation in the cardenolide content, thin-layer chromatography profiles, 
and emetic potency of monarch butterflies, Danaus plexippus reared on the milk- 
weed, Asclepias eriocarpa in California. J. Chem. Ecol. 8:579-633. 

COHEN, J. A. & L. P. BROWER. 1982. Oviposition and larval success of wild monarch 
butterflies (Lepidoptera: Danaidae) in relation to host plant size and cardenolide 
concentration. J. Kans. Entomol. Soc. 55:343-348. 

Dixon, C. A., J. M. ERICKSON, D. N. KELLETT & M. ROTHSCHILD. 1978. Some ad- 
aptations between Danaus plexippus and its food plant with notes on Danaus chry- 
sippus and Euploea core (Insecta: Lepidoptera). J. Zool., Lond. 185:437—467. 

ERICKSON, J. M. 1973. The utilization of various Asclepias species by larvae of the 
monarch butterfly Danaus plexippus. Psyche 80:230-244. 

FINK, L. S. & L. P. BROWER. 1981. Birds can overcome the cardenolide defense of 
monarch butterflies in Mexico. Nature 291:67-70. 

FINK, L. S., L. P. BROWER, R. B. WAIDE & P. R. SPITZER. 1983. Overwintering monarch 
butterflies as food for insectivorous birds in Mexico. Biotropica 15:151-153. 

MALCOLM, S. B., B. J. COCKRELL & L. P. BROWER. 1987. Monarch butterfly voltinism: 
Effects of temperature constraints at different latitudes. Oikos (in press). 

NIsHIO, S. 1980. The fates and adaptive significance of cardenolides sequestered by 
larvae of Danaus plexippus (L.) and Cycnia inopinatus (H. Edwards). Ph.D. Thesis, 
University of Georgia, Athens. 119 pp. 

NisHIo, S., M. S. BLUM & S. TAKAHASHI. 1983. Intraplant distribution of cardenolides 
in Asclepias humistrata (Asclepiadaceae), with additional notes on their fates in 
Tetraopes melanurus (Coleoptera: Cerambycidae) and Rhyssomatus lineaticollis 
(Coleoptera: Curculionidae). Mem. Coll. Agric., Kyoto Univ. 122:43-52. 

PRICE, P. W. & M. F. WILLSON. 1976. Some consequences for a parasitic herbivore, 
the milkweed longhorn beetle, Tetraopes tetrophthalmus, of a host-plant shift from 
Asclepias syriaca to A. verticillata. Oecologia 25:331-340. 

1979. Abundance of herbivores on six milkweed species in Iilinois. Am. Midl. 
Nat. 101:76-86. 

RAWLINS, J. E. & R. C. LEDERHOUSE. 1981. Developmental influence of thermal be- 
havior on monarch caterpillars (Danaus plexippus): An adaptation for migration 
(Lepidoptera: Nymphalidae: Danainae). J. Kans. Entomol. Soc. 54:387—408. 

ROESKE, C. N., J. N. SEIBER, L. P. BROWER & C. M. MOFFITT. 1976. Milkweed car- 
denolides and their comparative processing by monarch butterflies (Danaus plex- 
ippus L.). Rec. Adv. Phytochem. 10:93-167. 

ROTHSCHILD, M., J. VON Euw, T. REICHSTEIN, D. A. S. SMITH & J. PIERRE. 1975. 
Cardenolide storage in Danaus chrysippus (L.) with additional notes on D. plexippus 
(L.). Proc. R. Soc. Lond. (B) 190:1-31. 

SCHROEDER, L. A. 1976. Energy, matter and nitrogen utilization by the larvae of the 
monarch butterfly Danaus plexippus. Oikos 27:259-264. 

URQUHART, F. A. 1960. The monarch butterfly. University of Toronto Press, Toronto. 
361 pp. 

Wixsur, H. M. 1976. Life history evolution in seven milkweeds of the genus Asclepias. 
J. Ecol. 64:223-240. 

Woopson, R. E. 1954. The North American species of Asclepias L. Ann. Mo. Bot. 
Gard. 41:1-211. 

ZALUCKI, M. P. & R. L. KITCHING. 1982. Dynamics of oviposition in Danaus plexippus 
(Insecta: Lepidoptera) on milkweed, Asclepias spp. J. Zool., Lond. 198:103-116. 





Journal of the Lepidopterists’ Society 
40(4), 1986, 264-270 


BIOLOGY AND IMMATURE STAGES OF 
CITHERONIA SPLENDENS SINALOENSIS AND 
EACLES OSLARI IN ARIZONA (SATURNIIDAE) 


PAUL M. TUSKES 
7900 Cambridge 111D, Houston, Texas 77054 


- ABSTRACT. Citheronia splendens sinaloensis and Eacles oslari occur in Cochise, 
Pima, and Santa Cruz counties in southern Arizona. Both species have one generation 
per year. The flight season of E. oslari extends from early June to mid-August, and the 
larval host plants include Quercus species. The flight season of C. splendens extends 
from July to mid-August, and the larval host plants include wild cotton, manzanita, and 
New Mexico evergreen sumac. The immature stages are described for the first time. 


The citheroniine fauna of Arizona is unique in that all seven species 
are primarily of Mexican origin (Tuskes 1985). The biology of the two 
largest species, Citheronia splendens sinaloensis (Hoffmann) and Ea- 
cles oslari Rothschild, is poorly known. Ferguson (1971) illustrated the 
adults, summarized existing information, and indicated that their im- 
mature stages were undescribed. The purpose of this paper is to de- 
scribe the immature stages of both species and to present additional 
biological and distributional information. 


Citheronia splendens sinaloensis 
(Figs. 1-4) 


Citheronia splendens sinaloensis is the only member of the genus 
presently known to occur in Arizona. Citheronia mexicana G. & R. 
occurs just south of Arizona, in Sonora, Mexico. Although reported 
from Arizona before the turn of the century, there are no recent United 
States records. Citheronia regalis (F.) and C. sepulcralis (Druce) are 
common in the eastern or central United States but do not occur farther 
west than central Texas. 

Until recently splendens was only known from a few locations in 
Arizona, and specimens were scarce (Ferguson 1971). Observations and 
collecting in southern Arizona during the past 15 years have improved 
our knowledge of this species. Recently, splendens was taken from the 
Baboquivari Mts. (Pima Co.) east through Santa Cruz and southern 
Cochise counties to Guadalupe Canyon in the Peloncillo Mts. along the 
Arizona—New Mexico border, a distance of approximately 250 km. 

Adults were taken at lights from the first or second week of July to 
the second week of August, with peak emergence occurring in late 
July. Males appear at lights usually after 2300 h, and are collected 
sporadically until predawn. Females are seldom attracted to lights. 
Mating occurs between 0100 and 0330 h. 


VOLUME 40, NUMBER 4 265 


Males exhibit little variation in forewing pattern, except in the extent 
of the cream colored markings through the antemedial area. The 
hindwings show more variation than the forewings. Some specimens 
exhibit diffused white patches through the dark gray submarginal area. 
The basal area of the hindwings can be cream or dark gray. Male 
forewing length varies from 48 to 56 mm, averaging 53 mm (N = 21). 
Females are larger than males, with forewing length ranging from 59 
to 70 mm, averaging 64 mm (N = 12). Although forewing markings 
are similar to those of males, the wings of females are broader and less 
pointed, and the cream colored markings on the hindwings are reduced 
to small patches. 

Adults and larvae are usually associated with high desert arroyos 
containing wild cotton (Gossypium thurberi L.) (Malvaceae), and tran- 
sitional areas where manzanita (Arctostaphylos pungens H.B.K.) (Er- 
icaceae), and New Mexico evergreen sumac (Rhus choriophylla Woot. 
& Standl.) (Anacardiaceae) grow. Arizona black walnut (Juglans major 
Torr.) (Juglandaceae) is associated with canyon washes above 670 m, 
and is a possible but unconfirmed natural host. Wild cotton and man- 
zanita are the two most commonly utilized host plants. Some popula- 
tions are highly host specific and utilize only one species even though 
others are present. Ova are deposited on the underside of the leaves, 
either singly or in groups up to four. 

On cotton, the larva rests on the upper surface of the leaf curled in 
a J shape, and appears similar to a bird dropping (Fig. 2). During the 
early instars, most feeding occurs in the evening. As the larva matures, 
it rests on the petiole of the leaf or stem, and feeds sporadically through 
the day. Once larvae become established on the host plant, they are 
often reluctant to feed on other host plant species. Larvae transferred 
from cotton to manzanita, or from California pepper (Shinus molle 
L.) to cotton, often refuse to feed or fail to achieve the size of wild 
specimens. There are five instars, and development is rapid, with pu- 
pation occurring in September. Before pupation the larva leaves the 
plant and burrows into the ground where it constructs a pupation 
chamber of soil. 


Larval Description 


The following larval description is based on material reared from 
ova deposited by a female collected at Pena Blanca Lake, Santa Cruz 
Co., Arizona. Larvae collected from Box Canyon (Pima Co.) were also 
examined. Twenty-two larvae were examined; preserved larvae are in 
the author’s collection. 


First instar. Head: Black, diameter 1.5 mm. Body: Ground color black with orange 
band (Fig. 3). Length 15 mm, width 2.2 mm. Dorsal and dorsolateral thoracic (T) scoli 


266 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-6. 1-4. Immature stages of Citheronia splendens sinaloensis. 1, Mature fifth 
instar; 2, First instar in bird dropping pose; 3, First instar; 4, Third instar. 5,6. Immature 
stages of Eacles oslari. 5, Mature fifth instar; 6, First instar. 


black with bulb at tip, and 3-4 larger than dorsal abdominal scoli. Mid-dorsal caudal 
scolus on abdominal (A) segment 8 enlarged compared to dorsal abdominal scoli. All scoli 
black with short black spines on shafts. Segments A2, 6-8 with orange area on lateral 
surface surrounding each dorsolateral scolus. Segments A3-5 with orange extending from 
lateral surface dorsally over back; mid-dorsal black dot and smaller intersegmental black 
dot anterior to dorsal and dorsolateral scoli. True legs and prolegs black. 

Second instar. Head: Black, diameter 2.1 mm. Body: Ground color black and orange. 
Length 24-26 mm, width 5 mm. Dorsal T2, 3 scoli 5x or greater in length than dorsal 
abdominal scoli. Mid-dorsal scolus (A8) 8x or greater in length than dorsal abdominal 
scoli. All scoli black with black spines on shaft. Segments Al—8: Lateral surface from just 
below lateral scoli to base of dorsal scoli orange. Orange coloration extending over mid- 


VOLUME 40, NUMBER 4 207 


dorsal area on AS-5, as in first instar, but more variable. Caudal segment, true legs, 
prolegs, and spiracles black. 

Third instar. Head: Shiny black, diameter 2.5 mm. Clypeus black. Body: Ground 
color black or dark brown and orange (Fig. 4). Length 28-31 mm, width 6.7 mm. Dorsal 
thoracic and mid-dorsal scolus 2x or greater in length than abdominal scoli. All scoli 
black with short black spines on shaft. Thoracic segments primarily black. Abdominal 
segments: Area from base on lateral to dorsolateral scoli orangish brown. A series of black 
lines occurs on the lateral surface. Segments Al-8 with black line extending from just 
ventral of spiracle dorsally at 45-degree angle, terminating in intersegmental area. Seg- 
ments A3—5: Orangish brown coloration extends dorsally over back. Circular black patch 
occurs between dorsal and dorsolateral scoli. Segments Al, 2, and 6—9 with black to dark 
brownish orange dorsal area. Segments Al-8 with 1-2 black lines extending from mid- 
dorsal area ventrally, terminating posteriorly to spiracle. Intersegmental area dark brown 
with black triangular patch in line with dorsal scoli. Ventral surface orangish brown, 
with numerous parallel black lines in intersegmental folds. True legs and prolegs black 
and orange. Spiracles black. 

Fourth instar. Head: Black except for light brown frontal, frons, and adfrontal areas, 
diameter 4.6 mm. Clypeus cream. Body: Ground color dark purple to purplish pink. 
Length 59-66 mm, width 9.5-11 mm. Scoli of T2, 8 recurved, dorsal and dorsolateral 
10.5 and 5.5 mm long, respectively. Dorsal scoli T1 straight, 7.5 mm long, shaft white 
with black tip; short black spines on shaft. Mid-dorsal A8 scolus elongated, 10.5 mm long, 
with black shaft and short black spines. Dorsal, dorsolateral, and lateral scoli black with 
short black spines on shaft. Base of all abdominal scoli orangish brown. Abdominal 
segments: Dorsal, dorsolateral, and lateral surfaces dark purple to purplish pink. Posterior 
and anterior portion of abdominal segments with 1-2 thin black vertical lines. Small 
black vertical line and 2 black dots occur between dorsal and dorsolateral scoli. Thoracic 
segments primarily black dorsally, lateral surface black to dark purple. Ventral surface 
purplish brown; intersegmental area with 1-2 thin parallel black lines. True legs black 
to light brown. Proleg lateral surfaces black to dark purple, anterior and posterior surfaces 
light brown. Spiracles black. 

Fifth instar. Head: Shiny black, frontal suture brown, frons light brown, diameter 9.2 
mm. Clypeus light brown. Bedy: Ground color pale purplish brown. Length 110-117 
mm, width 15-17 mm. Dorsal and dorsolateral thoracic scoli T2, 3, and mid-dorsal scolus 
A8, 2x or greater in length than dorsal abdominal scoli. All dorsal, dorsolateral and 
lateral scoli light orange to light brown with black tips and small black spines on shaft. 
Sublateral scoli on T2, 3, and A7-9 light brown. Abdominal segments pale purplish 
brown. Intersegmental area light brown with thin black vertical lines. Cream colored 
undulating subspiracular fold extending from T8 to A8. Numerous thin black lines or 
dots on dorsal and lateral surfaces. One small black dot occurs anterior to dorsolateral 
scoli, and two between dorsal and dorsolateral scoli. T3-A8 with V-shaped mid-dorsal 
marking on each segment. Anterior to spiracle, black dots form continuous line to mid- 
dorsal area. Thin black line paralleling intersegmental fold. Ventral surface brownish 
black; intersegmental area with 1-2 thin parallel black lines. True legs light brown. Proleg 
lateral surface dark purple, anterior and posterior surfaces light brown. Spiracles black. 


Mature larvae of U.S. Citheronia are readily distinguished from 
each other. Larvae of splendens (Fig. 1) have a purplish brown ground 
color. Dorsal and dorsolateral thoracic scoli are light brown at the base 
with black tips. There is a trace of a cream colored abdominal subspi- 
racular line, and the ventral surface is brownish black. Citheronia 
mexicana occurs in Sonora, Mexico and has been reported from Ari- 
zona. Mature larvae of mexicana have a dark brown to black ground 
color and a prominent light yellow subspiracular line that extends the 
length of the abdomen. The dorsal and dorsolateral thoracic scoli are 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


light pink with black tips. Larvae of regalis have a brownish green 
ground color and bright red and black dorsal thoracic scoli. Bold white 
spiracular patches form a line on the lateral abdominal surface, but no 
subspiracular line is present. The ventral surface is greenish. Larvae of 
sepulcralis have a brown ground color and light brown dorsal and 
dorsolateral thoracic scoli. No spiracular or subspiracular white or cream 
colored lines or patches are present on the lateral surface, and the 
ventral surface is brown. Neither regalis or sepulcralis occur west of 
central Texas and are thus allopatric relative to splendens. 


Eacles oslari 
(Figs. 5, 6) 


Eacles from southern Arizona was assumed to represent a disjunct 
population of Eacles imperialis (Drury) until Ferguson (1971) elevated 
it to species status, based on differences in distribution, adult mor- 
phology, coloration, and pattern. At that time, E. oslari was rare in 
collections, and only 16 specimens were available for examination, all 
of which were from the Nogales—Pena Blanca Lake area of Santa Cruz 
Co., Arizona. Since then, oslari has been found in lower Madera Can- 
yon and Box Canyon in southern Pima Co., and in Patagonia, and most 
of the major canyons in the Huachuca Mts. of western Cochise Co. It 
is also widespread in Sonora and Sinaloa, Mexico. Although previously 
unreported from Mexico, oslari appears to be primarily a Mexican 
species whose northern limits extend into southern Arizona. 

The flight season of oslari in Arizona extends from the first week of 
July to mid-August, but the peak flight period is from 20 July to 5 
August. Both sexes are attracted to lights, but males are more frequent- 
ly captured than females. Forewing length of males ranges from 51 to 
58 mm, averaging 56 mm (N = 27); females are larger, forewing 
lengths ranging from 64 to 68 mm, averaging 66 mm (N = 88). There 
are three distinct adult color phenotypes resulting from differences in 
ground color: (1) deep yellow, (2) orangish brown, or (3) pale lavender 
to lavender-tinged. Some specimens appear transitional; yellow is the 
most common phenotype. 

In captivity, females deposit ova singly or in clusters of up to six. 
Confirmed larval food plants are Quercus oblongifolia Torr. Mexican 
blue oak, and Q. emoryi Torr., Emory oak, but other species of oak 
are probably utilized. Early instars are brown; after the third instar, 
larvae may be either brown or green. Before pupation, the larva leaves 
the tree and constructs a pupation chamber underground. In captivity, 
larvae have been successfully reared on various species of oak, Cali- 
fornia pepper tree, Schinus molle L. (Anacardiaceae), and Liquid- 
ambar styraciflua L. (Falatingiaceae). There is one generation per year. 


VOLUME 40, NUMBER 4 269 


Pine, a common host for E. imperialis nobilis Neumoegen, E. impe- 
rialis pini Michener, and some southern populations of nominate im- 
perialis, appears unsuitable as a host for oslari. An attempt was made 
by Gage (1976) to rear 40 oslari from ova on various conifers, but only 
one larva reached the last instar. Although the larva was illustrated in 
color, the immature stages were not described. 

Hybridization studies conducted by the author support the species 
rank assigned to oslari based on morphological criteria (Ferguson 1971). 
A newly emerged imperialis nobilis female, reared from ova collected 
in Waller Co., Texas, was tied out in Box Canyon, Pima Co., Arizona. 
The female attracted and mated with a male oslari at 0130 h, and the 
pair remained together until the following evening. Only 21 ova were 
deposited by the female. Seventeen were infertile, four developed em- 
bryos, but only one hatched, and it perished in the first instar. 


Larval Description 


The larval description is based on material reared from ova depos- 
ited by a female collected at Pena Blanca Lake, Santa Cruz Co., Ari- 
zona. Larvae reared by the author from females captured at Patagonia, 
Cochise Co. (N = 14) and Lower Madera Canyon, Pima Co. (N = 22) 
were also examined and are in the author’s collection. 


First instar. Head: Yellow with short light brown setae present, diameter 1.8 mm. 
Body: Ground color brown (Fig. 6). Length 11-13 mm, width 2.7 mm. Dorsal abdominal 
scoli black with small spines on shaft and one on apex. Dorsolateral and lateral scoli black 
with 1-2 black spines on shaft. Sublateral scoli on T1-2 and A6-7 consisting of short, 
simple black spines. Thoracic segments with enlarged dorsal and dorsolateral scoli which 
have forked tips. Mid-dorsal caudal scolus on A8 black with forked tip, at least 3x larger 
than dorsal abdominal scoli. Abdominal segments with 2 dark brown lines extending 
from lateral surface dorsally over back. Small crescent shaped line occurs anterior and 
ventral of spiracle. Ventral surface brown. True legs dark brown to black. Prolegs brown 
with black shields. 

Second instar. Head: Diameter 2.7-3.0 mm. Black, frontal area near mandibles or- 
ange. Short black secondary setae present. Diameter 2.7-3.0 mm. Body: Ground color 
dark brown. Length 20 mm, width 4.5 mm. Dorsal, dorsolateral, and lateral abdominal 
scoli cream to brown with 1—4 black spines. Enlarged mid-dorsal caudal scolus with light 
brown shaft and numerous short black spines. Dorsal and dorsolateral thoracic scoli 
orangish brown to light brown with numerous short black spines on shaft; 2 black spines 
at tip of each scoli. Lateral abdominal surface dark brown with numerous light brown 
secondary setae. Thoracic segments dark brown. Ventral surface brown. True legs or- 
angish brown. Prolegs brown with black shields. Spiracles black. 

Third instar. Head: Brown, area adjacent to adfrontal region light brown. Short brown 
secondary setae present. Diameter 3.7-3.9 mm. Body: Ground color brown or green. 
Length 25-28 mm, width 6.2 mm. Enlarged dorsal and dorsolateral thoracic scoli red 
with short red spines. All abdominal scoli cream; short cream spines with black tips on 
shafts. Enlarged caudal scolus red with red spines on shaft. Anal shield light brown with 
bluish black center; numerous small raised circular blue or white spots present. Green 
form: Spiracles turquoise and ringed with black. Proleg green; shields brown. True legs 
light brown. Brown form: Spiracles black and ringed by an inner gray and outer black 
line. Prolegs dark brown; black shields. True legs light brown. Both color forms with 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


elongate setae present on dorsal and dorsolateral areas, short secondary setae on lateral 
surface and prolegs. ; ; eae or . 

Fourth instar. Head: Dark brown or green, depending on body ground color. Light 
brown line extending from antennae, tapering dorsally to vertex of each lobe. Diameter 
5.5 mm. Body: Ground color brown or green. Length 41-51 mm, width 10 mm. Brown 
form: Enlarged dorsal and dorsolateral thoracic scoli brownish purple, with short yellow 
spines on shaft. Dorsal abdominal scoli yellow with short black spines. Dorsal, dorsolat- 
eral, lateral, and sublateral scoli light yellow and greatly reduced in size. Mid-dorsal 
caudal scolus brownish purple and enlarged. True legs light brown. Prolegs brown. Green 
form: Ground color green. Other coloration as above with following exceptions. Enlarged 
dorsal and dorsolateral thoracic scoli yellowish. True legs yellowish. Prolegs green. Both 
forms with elongated setae on dorsal and dorsolateral area, short secondary setae on 
lateral surface and prolegs. 

Fifth instar. Head: Brown or green depending on body ground color. Light brown 
line extending from antennae, tapering dorsally to vertex of each lobe. Diameter 7.8 
mm. Body: Ground color brown or green (Fig. 5). Length 90-110 mm, width 18 mm. 
Brown form: Dorsal and dorsolateral meso- and metathoracic scoli enlarged, 2.5 mm 
long, pinkish beige with small yellow tubercles on shaft. Dorsal, dorsolateral, lateral, and 
sublateral abdominal scoli reduced, 1 mm long, ivory in color with 2—4 short spines. 
Enlarged mid-dorsal caudal scolus pinkish beige with small yellow tubercles on shaft. 
Ventral and lateral surface brown, dorsolateral and dorsal areas light brown to brownish 
pink. Green form: As above but ground color green. Both color forms with elongate 
setae 9-11 mm long on dorsal surface. Lateral and ventral surfaces with light brown to 
short white secondary setae. Prothoracic shield yellowish green, ringed with light brown 
tubercles. Anal shield dark blue to black with small, raised light blue or white circular 
spots; shield ringed with light orange; caudal proleg shield similar. True legs yellowish 
brown. Spiracles turquoise. 


Larvae of oslari are similar in appearance and size to those of im- 
perialis from the eastern U.S. Larval variation among imperialis pop- 
ulations (Michigan, Virginia, Florida, Mississippi, and Texas) is so ex- 
tensive that characters which appear to make oslari larvae unique are 
also found in imperialis. The larvae of oslari tend to have less con- 
trasting differences in ground color on the dorsal and lateral surfaces, 
and the spiracles are consistently turquoise. Most populations of im- 
perialis have cream colored spiracles—others black or turquoise—and 
contrasting light brown patches on the lateral surfaces. 


ACKNOWLEDGMENTS 


I thank Noel McFarland, Michael Collins, and James Tuttle for comments on the 
manuscript, and James S. McElfresh for assistance in the field. 


LITERATURE CITED 


FERGUSON, D. C. 1971. The moths of America north of Mexico. Fascicle 20.2a Bom- 
bycoidea (in part). Classey, London. Pp. 1-174. 

GaGE, E. V. 1976. The immature stages of Eacles oslari (Citheroniidae). J. Res. Lepid. 
15:175-176. 

TUsKES, P. M. 1985. The biology and immature stages of Automeris randa and Au- 
tomeris iris hesselorum. J. Lepid. Soc. 39:163-170. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 271-288 


NEW GENERA FOR THE NEOTROPICAL “PAREROMENE™ 
SPECIES (PYRALIDAE: CRAMBINAE) 


DAVID E. GASKIN 


Department of Zoology, University of Guelph, 
Guelph, Ontario, Canada NIG 2W1 


ABSTRACT. Three new genera of neotropical diptychophorine Crambinae, Cleo- 
eromene, Neoeromene and Incaeromene, are established to contain seven species hitherto 
listed under Pareromene Osthelder (a synonym of the Old World genus Glaucocharis 
Meyrick), and two new species. Pareromene smithi (Druce) is transferred to Cleoeromene. 
Pareromene felix (Meyrick), P. herstanella (Schaus), P. octavianella (Zeller), P. parvalis 
(Walker), and P. straminella (Zeller) are transferred to Neoeromene, while N. parvi- 
puncta from Brazil is described as new. Pareromene excitata (Meyrick) was concluded 
to be synonymous with P. parvalis, and P. leucanthes (Meyrick) with P. felix. Incaeromene 
accommodates the single species subuncusella described as new. Two species groups are 
recognized within Neoeromene and their relations are briefly discussed. 


Seven neotropical species of diptychophorine Crambinae which were 
correctly removed from Diptychophora Zeller by Bkeszynski (1967) are 
still listed in Pareromene Osthelder. They cannot be retained under 
this name because it is synonymous with Glaucocharis Meyrick, a ge- 
nus confined to the Old World (Gaskin 1985). Furthermore, all differ 
from Glaucocharis in significant morphological characters. While the 
whole group probably had a common origin, there is no question of 
the neotropical forms being congeneric with either Glaucocharis or 
Diptychophora. The latter genus was recently revised (Gaskin 1986). 

Objectives of the present paper are to establish three new genera 
representing the lines of evolutionary divergence evident in the mor- 
phology of these neotropical forms, to describe two new species, to give 
two new synonymies, to illustrate characters not previously published 
for the known species, and to summarize features that distinguish the 
new genera from other genera of diptychophorine Crambinae. 

If experience with Old World diptychophorines is any guide, neo- 
tropical genera of these small and inconspicuous moths are probably 
undercollected at present. Cladograms based on present knowledge 
probably would not fairly represent the true diversity of existing species. 
Nevertheless, on balance of apparent apomorphies, Cleoeromene and 
Incaeromene represent derivative lines from Neoeromene-like stock 
with unspecialized and specialized valvae, respectively. Within Neo- 
eromene itself, two sets of trends can be recognized in male genitalia. 
The parvalis group (parvalis, herstanella, octavianella), with apically 
cleft uncus, strongly developed segregation of the sacculus, and aedea- 
gus cleft apically into a pair of strong protrusions, seems to be more 
derivative than the felix group (felix, straminella), where the uncus is 


PAPA JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


entire, the sacculus only weakly segregated, and the aedeagus has simple 
apical sclerotization. I do not separate these lines into named genera 
because parvipuncta, with a slight medial depression at the apex of the 
uncus, a partially segregated lobe of the sacculus, and paired patches 
of apical sclerotization of the aedeagus not drawn out into actual pro- 
trusions, seems to provide a clear link between the two group-trends 
genitalically. 

The following abbreviations are used in the text: BMNH (British 
Museum [Natural History]), CM (Carnegie Museum of Natural History, 
Pittsburgh), CNC (Canadian National Insect Collection, Ottawa), MCU 
(Museum of Cornell University), MNHU (Museum fiir Naturkunde der 
Humboldt-Universitat, Berlin, German Democratic Republic) and GC 
(Private collection of the author). Specimens examined bear Gaskin 
and Shaffer determination labels (BMNH). In descriptions of male gen- 
italia the LMB ratio refers to the length to median breadth ratio of the 
aedeagus. Decimals indicate position as a proportion of the total length 
of a structure or organ. In the forewing, measurements along the costa 
are taken from the base, those along the termen or margin from the 
apex, and those along fascia from the costa. In male genitalia, mea- 
surements along the uncus, gnathos, valva, valval costa and aedeagus 
are from the base of each. In female genitalia, measurements along 
the ductus bursae are from the ostium. 


Key to the Genera of New World Diptychophorine Crambinae 


1 Forewing vein R, vestigial or absent, hindwing M, absent (both sexes). Lateral, 


medial foramen present in vinculum of male oc ecceeeeeeeeneen Microcausta 
- Forewing vein R, and hindwing M, fully developed. Male vinculum without 
FOTAIMCD acicccececennesrtgenninnetncdoned to Ue 2 
2(1) 6: Gnathos strongly “‘fish-hooked”; valva quadrate, truncated, barely as long as 
wide. 
9: Antrunmiarmenn brati@trsey Sinn ley mite eee anes ae se Diptychophora 


- 6: Gnathos slightly curved or nearly straight. Valva about 1.5x longer than 
wide, often tapered apically. 


9: Antrum) sclerotized, often.complex 2.) 2 ee 3 
3(2) 6: Juxta with two pairs of sclerotized apical horns. 
¢: Antrum with complex folds but not segregated into laterally paired structures 
cal ok CN See NN se 1 Oe Ee Steneromene 
— 6: Juxta simple to complex, but without two pairs of protrusions. 
¢: Antrum complex, with strong tendency for division into right and left lateral 
structures, including subantral accessory sacs... 4 
4(3) 6: Juxta with huge medial spur. Gnathos hastate. Aedeagus lacking sclerotiza- 
tion. 
¢: Antrum with pair of internal lateral cupped folds. Corpus bursae with two 
sigma 20 kd SCY Pea ea ad Cleoeromene 


— 6: Juxta lacking spur. Gnathos not hastate. Aedeagus with some apical scleroti- 
zation, often laterally divided. 

: Antrum with a pair of internal lateral folds, but not cupped. Corpus bursae 
asignate: 08 ce ee Na 2) ee 5 


+0 


VOLUME 40, NUMBER 4 Did 


5(4) 6: Tegumen simple, uncus without basal spurs. 


So ANGIE AG RRs eg ea Ae 2 et SA ROS RNR © tel RIE A RNA le Neoeromene 
— 6: Tegumen with prominent apical posteriad spur. Uncus with 2 dorsal spurs. 
ermal ese ee kn wil seme te 2 ee Bs ae Incaeromene 


Cleoeromene, new genus 


Type species Diptychophora smithi Druce (1896:292, pl. Ixiv, fig. 20) (by monotypy). 


Description. Forewing Sc concurrent with R,, R, and R, stalked. Hindwing cell nearly 
closed by connections between M,, M,; M, and Cu, arising from cell, but with roots close 
together. Male juxta bearing at its ventroposterior center a gigantic apically serrated, 
flattened prong. Valva tapering, with subapical prong arising from distal extremity of a 
saccular fold, but involved with costal region by introrse movement. Female antrum 
characterized by a pair of large, strong, lateral, internal, elongate cupped folds. Corpus 
bursae with two small circular signa. 

Etymology. K)éoo (Kleos)—glory (Doric Greek); Epwuéevn (Eromené)—mistress (f). 


Cleoeromene smithi (Druce), new combination 


Diptychophora smithi Druce (1896:292). 
Diptychophora smithi Druce; Bleszynski & Collins (1962:299). 
Pareromene smithi (Druce) Bleszynski (1967:92, 96). 


Description (Fig. 1). Wing expanse 15-17 mm (N = 11). External features satisfactorily 
described by Druce (1896). 

Male genitalia (Fig. 9) (N = 1). Uncus simple, swollen dorsally from 0.3-0.9. Gnathos 
tapering abruptly to slender pedicel, with spatulate-hastate, flattened apical expansion. 
Tegumen simple, with strong ventral margins. Vinculum narrow, barely half as wide in 
profile as length of uncus. Saccus narrow, elongate, almost tubular with rounded apex; 
equal in length to uncus. Juxta a quadrate plate, basally triangular, rounded apically, 
with concave lateral margins. A huge prong arising from upper central region of juxta, 
grotesquely out of proportion to rest of genitalia, nearly 4x length of uncus, is apparently 
formed of two fused elements and probably anellar or transtillar in origin. Valva rounded, 
tapered, with some centrobasal development which cannot be attributed for certain to 
development of the dorsal margin of the sacculus or ventral margin of the valvula region. 
Costa strong, drawn apically into short, blunt prong. Valvula with longitudinal medial 
pleat. Aedeagus about 1.2 length of valva, tubular, LMB ratio about 10:1, apically 
truncate, lacking cornuti or external spines. 

Female genitalia (Fig. 10) (N = 1). Anal papillae weakly fused dorsally. Anterior 
apophyses about 0.6, and 8th tergite about 0.5 length of posterior apophyses. Antrum 
a wide, flared funnel. Lamella antevaginalis highly sclerotized, developed into a pair of 
lateral, elongate, cupped plates curved below ostium (possibly forming a receptacle for 
the juxtal spur of the male). Lamella postvaginalis forming a strong, single, dorsal plate. 
Ductus bursae about 2.5 length of posterior apophyses. Scobinate subantral sac present 
at about 0.3, ductus seminalis joining at about 0.7. Corpus bursae bearing two small, 
circular signa. 

Types. Lectotype é (designated here), MEXICO: Amula, Guerrero, 6,000 ft (1,829 m), 
“August”, Smith, BMNH, genit. prep. pyral. 15101. Paralectotypes: 5 6 with same data, 
2 6 similar but “September’’, all BMNH. 

Other material examined. MEXICO: 2 4, 1 9, Iguala, Guerrero, 2,400’ (731 m), 
8. VIII.1954, J. G. Chillcott, CNC, genit. prep. D.E.G. 1979/8. 


Discussion. Little is known about this species, other than that it 
inhabits moderate altitudes in SW Mexico. The relations to Neoero- 
mene in genitalic characters in both sexes are clear, except that in 
smithi it is the juxta, rather than the valva, which has undergone 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-8. Forewings of Cleoeromene, Neoeromene and Incaeromene species. 1, 
Cleoeromene smithi; 2, Neoeromene octavianella; 3, N. herstanella; 4, N. parvalis; 5, 
N. parvipuncta, 6, N. straminella; 7, N. felix; 8, Incaeromene subuncusella. 


sclerotized elaboration and differentiation. The antrum is also more 
complex in Cleoeromene than in Neoeromene. 


Neoeromene, new genus 


Type species. ?Isopteryx parvalis Walker (1865:1316). 


Description. Forewing Sc concurrent with R,; R, stalked with R,; venation similar to 
Glaucocharis Meyrick. The male genitalia of Neoeromene characteristically have one or 
more large, sclerotized areas or spurs at the apex of the aedeagus, unlike any found in 
species of Glaucocharis. In about half the known species, the sacculus of the valva is 


VOLUME 40, NUMBER 4 OTS 





Fic. 9. Male genitalia of Cleoeromene smithi lectotype, posterior aspect with left 
valva (left), aedeagus (right). Scale = 1 mm. 


strongly developed, with medial and ventral flanges or points or both. In the others, 
lateral or medial grooving of the valva extends from this region. In most species, the tip 
of the uncus is minutely or distinctly bifid. There is a strong tendency in all species for 
marked medial constriction of the juxta, for development of apical curved horns on the 
juxta, and the parallel development of an anellar structure around the aedeagus not 
found in Glaucocharis, but similar to that in some southern hemisphere Chilonini (Gaskin 
1973). Relatively few females of Neoeromene are known, but in these the antrum is 
usually much broader and more complexly folded than in Glaucocharis, and the ductus 
is also broad and either pleated or laterally lobate. While the forewing markings are 
superficially similar to those of many species of Glaucocharis, there are marked differ- 
ences and developments in the morphology of the genitalia which do not at all resemble 
those found in the Old World genus. Because females of several species are not known, 
the sexes are keyed separately. 
Etymology. Néoo (Neos)—young; Epwpéevn (Eromené)—mistress (f). 


Key to the Species of Male Neoeromene 


1 Costal region of valva strong or weak, but lacking basal protuberance; saccular 
region of valva differentiated into strong lobe terminating in a ventral prong 
— Costal region of valva with prong; sacculus undeveloped cee 4 
2(1) Sclerotization of sacculus confined to ventral] margin eee eneeneenene nen 3 
—  Sclerotization extending thickly at right angles into middle of valva from base 
ele be INT TRIAL HRT ek ASP Ny BIN CURL RR 2 a eh octavianella 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 10,11. Female genitalia of Cleoeromene and Neoeromene, ventral aspect. 10, 
Cleoeromene smithi; 11, Neoeromene parvalis. Scale = 1 mm. 


VOLUME 40, NUMBER 4 DAT 


3(2) Terminal prong of sacculus short, barely 0.15x length of uncus; sclerotized 


anellar structure present in association with juxta 00 parvalis 
- Terminal prong of sacculus long, 0.5 or more length of uncus; membranous 
anellar structure present in association with juxta 00 herstanella 
4(1) Costal prong smooth, arising near base, directed dorsoposteriad 5 
— Costal prong heavily spinose, arising at about 0.5 and curving introrse and 
WER! ee era ne. MRE ai ae ee se 8 2 parvipuncta 
5(4) Juxta with pair of strong apical horns. Aedeagus with single narrow, apical 
PRON -o QU SUR eC En east 2 Tet A SOU a LS es ee straminella 
— Juxta lacking apical horns. Aedeagus with large, flattened, sclerotized apical 
spur, with serrated edge and secondary hooked basal structure felix 


Key to the Species of Female Neoeromene 


1 Medial depression in posterior margin of 7th sternum; antrum a strong, globate 
funnel, ductus bursae with several small accessory pockets in subantral region; 
CORPESRMUTSACSIN ASSIV EC aeee eee shee ESR ta ek parvalis 

— Posterior margin of 7th sternum without medial depression; antrum strong, but 
not globate; ductus bursae without accessory pockets, or if present, associated 
with very elongate, narrow ductus bursae 2 

2(1) Ductus bursae less than length of 8th tergum at antrum, barely wider than 
apophyses at midpoint, elongate, and sclerotized only in antral region _.. 
NIN land in Se ame OS eat ie bias Ne sem, cba ig ss Re herstanella 

— Ductus broad, 2x length of 8th tergum at antrum, strong to about 0.4, with 
series of longitudinal pleats from subantral region to junction with ductus 
SEEMING spe I eed ED ha A felix 


Neoeromene parvalis (Walker), new combination 


?Isopteryx parvalis Walker (1865:1316). 

Diptychophora excitata Meyrick (1931:109). NEW SYNONYMY. 

Diptychophora excitata Meyrick; Bleszynski & Collins (1962:297). 

Pareromene excitata (Meyrick) Bleszynski (1967:92, 96). 

[PIsopteryx parvalis (Walker) Bleszynski & Collins (1962:296, 298), erroneously synon- 
ymized with Diptychophora azanalis (Walker).] 

Pareromene parvalis (Walker) Bleszynski (1967:92, 96). 


Description. Alar span 10-12 mm (Fig. 4) (N = 7). Redescribed here, since Walker’s 
account is ambiguous. Labial palpi, head, thorax, abdomen, dull white, with scattered 
brownish scaling. Ground color of forewings dull white. Basal fascia nearly obsolete, 
position marked only by broken patches of dark scales. Antemedial fascia more complete, 
but solid, irregular, dark brown. Faint orange reniform marking present. Costa with 
large patch of white from median line to 0.6, terminated by short oblique bar on costa. 
Region between basal and antemedial fascia, and all of discal area, filled with scattered 
scales of buff proximally, darker brown distally. Postmedial fascia cream, irregularly and 
narrowly bounded with dark brown. Terminal zone creamy white, with faint orange 
stripe along margin, within which is a row of 3-4 irregular black spots. Apical zone faint 
orange, with large central white area. Cilia pale brown with dark apices. Hindwings 
creamy white, cilia similar, but with dark bases in apical region. Ventral surfaces straw 
and mid-brown, with apical forewing markings repeated from dorsal surface. 

Male genitalia (Fig. 12) (N = 3). Uncus broad, simple, convexly tapered to a cleft 
apex. Gnathos slender, curved dorsad to bluntly pointed apex. Tegumen simple; vinculum 
a narrow strap at base of valva; saccus almost negligible; juxta basally triangular, sharply 
constricted distally into a narrow folded groove having membranous extensions with 
sclerotized margins, forming a weak anellar structure supporting aedeagus. Valva 2.7- 
3.0x length of uncus, apically broadly rounded; costal region a narrow sclerotized zone; 
sacculus a strong marginal lobe comprising basal % of valva, terminating in short, sharp 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 12. Male genitalia of Neoeromene parvalis, posterior aspect with left valva (left), 
aedeagus (right). Scale = 1 mm. 


spur at about 0.4; valva crossed by narrow membranous zone from base of this spur 
obliquely to base of costal margin. Aedeagus tubular, LMB ratio about 8:1, curved gently 
ventrad, apex bearing pair of broad curved spurs, one larger than the other. 

Female genitalia (Fig. 11) (N = 1). Anal papillae broad, prominent, moderately fused 
dorsally; 8th abdominal tergum narrow, barely as long as anal papillae; anterior and 
posterior apophyses approximately equal in length; posterior margin of 7th sternum 
medially indented, infolded. Antrum a broad, globate, sclerotized funnel, basally con- 
stricted at about 0.2; ductus bursae complexly folded, with 2 or more small, membranous, 
lateral, subantral pockets; position of ductus seminalis junction difficult to determine in 
preparation, apparently at about 0.5; corpus bursae asignate, massive. 

Types. Holotype parvalis 6, BRAZIL: “Ega (Brazil) 57-125”, BMNH, genitalia prep. 
BM pyral 7679. 

Holotype excitata 2, BRAZIL: Obidos, VIII.19, Parish, BMNH genitalia prep. BM 
pyral 7681. 

Other material examined. PERU: 4 6, Iquitos, VIII.1920, Cornell Univ. Exp. Lot 607, 
MCU, genitalia prep. Cornell #1 (M. Shaffer); 1 4, 1. VIII.1920 (same Cornell lot. no.), 
CNC, genitalia prep. 4382-SB. 


Discussion. The genitalia of both sexes demonstrate clearly the ma- 
jor characteristics of this genus; the male uncus is apically divided, and 
the sacculus of the valva strongly developed, while in the female the 
broad antrum is sclerotized, the subantral region of the ductus is com- 
plex, and the corpus bursae asignate. Nothing is known of the geo- 


VOLUME 40, NUMBER 4 279 





Fic. 18. Male genitalia of Neoeromene herstanella holotype, posterior aspect with 
left valva (left), aedeagus (right). Scale = 1 mm. 


graphic distribution and ecology, except that it appears to occur across 
the upper Amazon region and the flight period includes August. 


Neoeromene herstanella (Schaus), new combination 


Diptychophora herstanella Schaus (1922:132). 
Diptychophora herstanella Schaus; Bleszynski & Collins (1962:297). 
Pareromene herstanella (Schaus) Bleszynski (1967:92, 96). 


Description. Alar expanse 9 mm (Fig. 3) (N = 2). External features adequately de- 
scribed by Schaus (1922). 

Male genitalia (Fig. 13) (N = 1). Uncus broad, tapered, bluntly pointed; apex minutely 
cleft. Gnathos slender, tubular, distal half rugose or ““‘pimpled”. Tegumen simple, broad, 
strong; vinculum triangular in profile, about half as wide as uncus length; saccus small, 
truncate, with angles slightly pointed; juxta an elongate plate, broad at base, otherwise 
narrow, weak; membranous anellar structure present. Valva 4.0-4.2x length of uncus; 
costal region a sclerotized strip running dorsal length of valva, without prongs or protru- 
sions; valvula tapered to blunt dorsal point; sacculus with distinct lobe, drawn distally 
into a prong directed posterioventrad, 0.5 x length of uncus. Aedeagus about 0.7 x length 
of valva, stout. LMB ratio about 5.5:1, with pair of large, recurved apical horns; cornuti 
absent. 

Female genitalia (Fig. 15) (N = 1). Anterior and posterior apophyses approximately 
equal in length; anal papillae separate. Antrum a sclerotized tapered funnel, with basal 
swellings; ductus bursae about 4.5 length of posterior apophyses, slender; ductus sem- 
inalis joining at about 0.4; corpus bursae relatively small and asignate. 


280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 






A AIS 

Sera ninsrn 

"Ngan * 
uva@ <I 


4 


IS 





Fics. 14, 15. 14, Female genitalia of Neoeromene felix (leucanthes holotype); 15, 
N. herstanella; both ventral aspect. Scale = 1 mm. 


Types. Holotype 4, PANAMA: with 4 labels; “Porto Bello, Feb. 24. Pan.” (white), 
“Type No. 25535 U.S.N.M.” (orange), “slide SB 6 No. 4617” (pale blue), “Diptychophora 


herstanella type. Schs.”’ (white). 
Other material examined. 1 ?, COSTA RICA: Siguirres, Limon, 50 m, no date. BM 


16753. 


VOLUME 40, NUMBER 4 281 





Fic. 16. Male genitalia of Neoeromene octavianella, paralectotype, posterior aspect 
with left valva (left), aedeagus (right). Scale = 1 mm. 


Discussion. This species is presently known only from Panama and 
Costa Rica, with a flight period including February. It is close to parvalis 
in major genitalic characters, and may be a Pleistocene segregate from 
that species. 


Neoeromene octavianella (Zeller), new combination 


Diptychophora octavianella Zeller (1877:33, pl. 1, fig. 13). 
Diptychophora octavianella Zeller; Bleszynski & Collins (1962:298). 
Pareromene octavianella (Zeller) Bleszynski (1967:92, 96). 


Description. Alar span 12-14 mm (Fig. 2) (N = 83). Described here because Zeller’s 
account does not fully distinguish it from similar species. Female not yet collected. Head, 
thorax, palpi dull whitish with some grey and buff. Ground color of forewings off-white. 
Basal fascia represented by some blackish brown scales. Antemedial fascia more distinct, 
dull brown, slightly zigzagged. Faint yellow reniform mark present in disc, which has 
scattered buff scaling. Postmedial fascia a broad creamy band, zigzagged near apical 
angle and towards dorsum, edged thinly with dark brown. Terminal zone white, with 
2-3 small blackish spots on margin between 0.5-—0.8, surrounded by some yellow scales. 
Apical zone bright yellow, with suboval, central, shining white patch not touching costa. 
Small wedge of white scales present at apical extremity. Hindwings whitish. Cilia pale 
brown with darker tips. All ventral surfaces dull mid-brown, with apical and terminal 
markings repeated from dorsal surface. 

Male genitalia (Fig. 16) (N = 2). Uncus broad, slightly setulose, tapered, with bifid 
apex. Gnathos slightly shorter than uncus, T-shaped, with strong base nearly at right 
angles to subapically swollen perpendicular element. Tegumen simple, with thick, strong 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 17. Male genitalia of Neoeromene parvipuncta, holotype, posterior aspect with 
left valva (left), aedeagus (right). Scale = 1 mm. 


dorsal and ventral margins; vinculum indistinguishable from saccular lobe of valva; saccus 
short, broad, straplike and posteriorly truncate; juxta subtriangular, apically tapered, 
indented basally and developed apically into pair of short prongs. Valva nearly 3x uncus 
length, costal margin strong to the tapered apex; sacculus-vincular region developed into 
a distinct lobe, ventral margin strong, with zone of sclerotization running introrse into 
central part of valva, almost touching base of costa, thick, short, curved thorn at 0.5. 
Aedeagus about 0.8 length of valva, stout, LMB ratio about 7:1, lacking cornuti, with 
apical portion divided to form pair of strong, apically recurved prongs lateral to ductus 
ejaculatorius. 

Types. Lectotype 6 (here designated), PANAMA: “Chiriqui Ribbe’, Coll. Staudinger, 
“Diptychophora octavianella” [blue label]; “Origin” [pink label]; “Typus’’ [red label], 
“Praep. Gen. Nr6708” [Bleszynski]; “D2.845 Lectotype” [circular purple-bordered label]; 
“LECTOTYPE Diptychophora octavianella Zeller det. M. Shaffer. 1976”, MNHU. 

Paralectotypes, 2 6, data as above except one has basic data on white (not blue) label, 
both bear “Paralectotype” [circular blue-bordered labels]; one bears “Abdomen missing” 
[blue label]; both bear “PARALECTOTYPE Diptychophora octavianella Zeller det. M. 
Shaffer. 1976” [white labels]; one bears “73?” [faded paper], MNHU. 


Discussion. Neoeromene octavianella, presently known only from 
Panama, forms part of a closely related cluster of species including N. 
parvalis and N. herstanella. 


VOLUME 40, NUMBER 4 283 


Neoeromene parvipuncta, new species 


Description. Alar span 11 mm (Fig. 5) (N = 1). Labial palpi 1.2x head length, 
yellowish, with blackish tips. Head, thorax and abdomen yellowish with occasional patches 
of paler cream and dark brown scales. Tarsi of forelegs banded alternately with buff and 
dark brown. Ground color of forewings golden yellow; basal fascia obsolete, marked only 
by a few irregular patches of blackish brown; antemedial fascia dark brown, irregular, 
incomplete, narrow. Discal region of forewing overlain with pale cream scaling; central 
area with some black or brown patches, but no distinct reniform stigma. Postmedial 
fascia white, distinct, broad, sharply edged with dark brown. Some fine dark neural 
streaks extend from disc to termen. Apical region orange, with a central pear-shaped 
white zone, edged posteriorly with dark brown, some dark brown shading on costa. 
Terminal region golden yellow, subapical marginal indentation distinct; termen bearing 
5-6 indistinct blackish spots set in small areas of white scales, from 0.3 to 0.9. Hindwings 
white, clouded near apex with pale brown. All cilia pale brown with dark brown tips. 

Male genitalia (Fig. 17) (N = 1). Uncus curved, bluntly tapered, laterally setulose. 
Gnathos slightly longer than uncus, apically swollen and cupped. Tegumen with strong 
posterior margins; saccus simple, broad, apically rounded; vinculum broad, almost as 
wide in profile as uncus length; juxta broad, quadrate, drawn into pair of anellarlike 
dorsal projections which close around aedeagus. Valva about 2.2 length of uncus, blunt- 
ly tapered; costal region strong, terminating in a heavily spinose protrusion arising at 
0.5-0.6 and curving introrse, posteriad, then ventrad inside valva. Valva with slight 
dorsoventral constriction at about 0.5; sacculus a spinose lobe partially segregated from 
base of valvula. Aedeagus about 0.8 length of valva, tubular, straight, truncate, with 
dorsal and ventral zones of apical-subapical sclerotization, not drawn out into apical 
prongs; LMB ratio about 5.5-6.0:1; cornuti absent. 

Type. Holotype 6, BRAZIL: Curitiba, Parana, 920 m, -.X.1975, V. O. Becker, in V. 
O. Becker collection, specimen 4823. 


Discussion. Nothing is known of the biology of this species; the type 
specimen was taken in October. On genitalic characters it is closely 
related to N. straminella. 


Neoeromene straminella (Zeller), new combination 


Diptychophora straminella Zeller (1877:32, pl. I, fig. 12). 

Diptychophora straminiella Zeller; Hampson (1896 (1895):943) (misspelling of strami- 
nella). 

Diptychophora straminella Zeller; Bleszynski & Collins (1962:299). 

Pareromene straminella (Zeller) Bleszynski (1967:92, 96). 


Description. Alar span 14-15 mm (Fig. 6) (N = 3). Redescribed here; there is more 
variation than indicated by Zeller. Female not yet collected. Labial palpi about 0.75 x 
head length, yellow, with blackish apices and transverse striping. Head, thorax, abdomen 
creamy yellow with sprinkling of chocolate scales. Ground color of forewings pale yel- 
lowish orange, basal fascia nearly obsolete. Double, faint, reniform stigma present, to- 
gether with some faint interneural streaks in discal region. Postmedial fascia creamy 
yellow, thinly edged with dark brown. Terminal zone yellowish, clouded with dark 
brown, except distally, where the margin bears a row of 5 black spots set in a narrow 
strip of orange-ochreous. Apical zone bright orange-brown, with a central triangular 
white mark, banded thinly with black proximally and posteriorly. White wedge of scales 
at apical costal extremity. Cilia pale brown with darker apices. Ventral surfaces straw, 
with some repetition of dorsal apical and terminal markings. 

Male genitalia (Fig. 18) (N = 2). Uncus narrow, tapered, bluntly pointed, curved 
ventrad. Gnathos subtubular, swollen abruptly at 0.5, then tapering to bluntly pointed 
apex, curved dorsad. Tegumen simple, with strong ventral margins. Vinculum broad, 
about 0.75 as wide as uncus length. Saccus large, broad, rounded, about as long as 


284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 18. Male genitalia of Neoeromene straminella, posterior aspect with left valva 
(left), aedeagus (right). Scale = 1 mm. 


uncus. Juxta an apicomedially cleft quadrate plate, more rounded basally than apically, 
fused with an anellar structure bearing a pair of large, curved, strong apical horns, bases 
of which are partially fused laterally with basal extremities of valval costa. Valva about 
2.5 length of uncus, sacculus weakly developed, costa strong, bearing a curved basal 
prong nearly half uncus length; costal lobe extends almost to apex of valva, apex discrete 
and partly separate. Aedeagus about 1.4 length of valva, curved slightly dorsad, LMB 
ratio about 8:1. Cornuti absent, but apex bears an elongate, blunt thorn dorsally. 

Type. Holotype 6, BRAZIL: “N. Friburgo Bres.”, Staudinger (also bears labels “Ori- 
gin’, “Typus’, “7:?”), MNHU, genitalia prep. GS-603-SB. 

Other material examined. 2 6, BRAZIL: Petropolis, Walsingham, BMNH, BM pyral 
15093. 


Discussion. So far collected only in Brazil, a close relative of hersta- 
nella and parvalis. Flight period unknown. 


Neoeromene felix (Meyrick), new combination 


Diptychophora felix Meyrick (1931:108). 

Diptychophora leucanthes Meyrick (1931:108); Bleszynski & Collins (1962:297). NEW 
SYNONYMY. 

Diptychophora felix Meyrick; Bleszynski & Collins (1962:297). 

Pareromene felix (Meyrick) Bleszynski (1967:96). 

Pareromene leucanthes (Meyrick) Bleszynski (1967:92, 96). 


VOLUME 40, NUMBER 4 285 





Fic. 19. Male genitalia of Neoeromene felix, lectotype, posterior aspect of left valva 
(left), aedeagus (right). Scale = 1 mm. 


Description. Alar span 12-13 mm (Fig. 7) (N = 10). External features adequately 
described (twice) by Meyrick (1931). Characters described for leucanthes overlap those 
for felix. ; 

Male genitalia (Fig. 19) (N = 2). Uncus simple, tapered, lightly setulose, curved ven- 
trad, with blunt apex. Gnathos about 0.8 uncus length, curved slightly dorsad, tapered 
abruptly near base, then parallel-sided until near slightly hooked, pointed apex. Tegumen 
simple; vinculum broadly triangular, about 0.7 <x as wide as uncus length; saccus broadly 
pyramidal, about 0.5 uncus length; juxta a subtriangular plate, with a sclerotized in- 
verted “Y”’ strengthening dorsal apex and expanded ventrolateral margins. Valva broad, 
sacculus barely differentiated, costal region strong to about 0.5, with thick, simple, slightly 
curved basal prong 0.7 uncus length. Valva parallel-sided to about 0.7 from base, then 
tapering abruptly to a quadrate, setulose, apical lobe. Aedeagus about 0.7 x valval length, 
tubular, truncate, stout, LMB ratio about 6.5:1, apex bearing a broad, flattened, spade- 
shaped, strongly sclerotized spur with serrate margin, and a second much smaller hooked 
spur below exit of ductus ejaculatorius. A single large, irregular cornutus present sub- 
apically. 

Female genitalia (Fig. 14) (N = 2). Anal papillae moderately fused, anterior and 
posterior apophyses nearly equal; 8th tergum about 0.8 x length of posterior apophyses; 
7th sternum rounded and partly tapered posteriorly; sternum margin turned introrse, 
combining with lamella antevaginalis into a lodicular structure while lamella turns ex- 
trorse, forming a protruding lip. Lateral extremities form a pair of nearly circular cuplike 
structures directed ventrad. Lamella postvaginalis set deep in the broad antrum, medially 
broadly fenestrate. Ductus bursae about 3.5 x length of posterior apophyses, with ductus 
seminalis joining at about 0.9, close to corpus bursae; ductus bursae pleated from 0.4- 
0.6; corpus bursae asignate. 


286 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 20. Male genitalia of Incaeromene subuncusella, holotype, posterior aspect with 
left valva (left), aedeagus (right). Scale = 1 mm. 


Types. N. felix: Lectotype é (selected by S. Bleszynski and designated here). BRAZIL: 
Obidos, “8.19’, Parish BMNH, genitalia pre. BM pyral 7682. 

Paralectotypes, BRAZIL: 4 6, one lacking abdomen, data as above but “8 & 9.19”, 
BMNH;; 1 8, data as above, BMNH, and bearing label in Bleszynski’s writing “BM 2 
7684”, but unfortunately the slide is missing from the collection; 1 6, Santarem, “8.19” 
Parish, BMNH, BM pyral 7683. 

N. leucanthes: Holotype 2, PERU: Iquitos “3.20”, Parish, BMNH, genitalia prep. BM 
pyral. 7682. 

Other material examined. GUYANA: 1 ? (lacks hindwings, frenula, abdomen): Con- 
fluence of Orinoque & New Rivers, 20.viii—-20.ix.1937, Hudson, BMNH; ?BOLIVIA: 1 9, 
“Carn. Mus. Acc. 6473” (no other data), CM, genitalia prep. GS-5983-SB. 


Discussion. Variation in the proportions of brown and yellow cloud- 
ing in different specimens is considerable. Based on his criteria for 
distinguishing species, Meyrick’s (1931) conclusions were reasonable. 
This species is probably widespread in the tropical forests of the north- 
ern and central regions of South America. The flight period is known 
to include August and September. 


Incaeromene, new genus 


Type species Incaeromene subuncusella, new species (original designation). 


Description. Venation as in Neoeromene. Genitalia characterized by singular devel- 
opment of dorsal region of tegumen into a hood, projecting over uncus and gnathos, 
terminating in a strong spur posteriad, also by costal lobe of valva bearing a broad 
capitulate protrusion. Sacculus strongly developed as in Neoeromene, but posteriorly 
heavily spinose on inner surface. 

Etymology. Inca—pertaining to Incas; Epwyevn (Eromené)—mistress (f). 


VOLUME 40, NUMBER 4 287 


Incaeromene subuncusella, new species 


Description. Alar span 15 mm (Fig. 8) (N = 4). Head, labial palpi, thorax, abdomen 
silvery white; medial surfaces of palpi and patagia with scattered chocolate-brown scaling. 
Ground color of forewings silvery white. Basal and antemedial fascia obsolete, positions 
marked by some brown scaling. Postmedial fascia composed of pair of curved brown 
lines. Termen with weak subapical indentation at about 0.25, and 3-4 black marginal 
spots from 0.5-0.9. Faint reniform mark present in disc amid patch of brown scaling, 
some similar clouding on costa at base and subterminal region. Cilia white with brown 
bars. Hindwings pure silvery white, as are cilia. Underside of forewings dull brown. 

Male genitalia (Fig. 20) (N = 1). Uncus and gnathos stout, tapered, pointed, slightly 
curved, setulose. Uncus with pair of sclerotized basal dorsal protrusions. Tegumen strong, 
drawn dorsoposteriorly into long hood over uncus, terminating in sharp, sclerotized 
point. Saccus negligible, vinculum narrow. Juxta weak, laterally folded and posteriorly 
drawn into an anellar structure around aedeagus. Valva about 2.7 uncus length, with 
broad sacculus, lobate posteriorly, heavily spinose. Costal region drawn into broad, blunt- 
ly capitulate setulose lobe. Aedeagus 2.2 uncus length, massive, apically lightly scler- 
otized, with pair of small dorsal subapical thorns. 

Type. Holotype 6, PERU: Cuzco, Pillahuata, 2600 m, 14-18. viii.1982, M. Matthews & 
M. Packer. BMNH genitalia prep. pyral. 17131, in BMNH. Paratypes 3 6, same data as 
holotype, all in BMNH. 


Discussion. This undistinguished looking species exhibits the most 
peculiar modification of the dorsal tegumen of any known diptycho- 
phorine. Without female genitalia, it is impossible to speculate about 
the function of this spurred hood over the uncus. 


ACKNOWLEDGMENTS 


I thank the Department of Entomology, British Museum (Natural History) for provid- 
ing space, facilities, and assistance, and the Biosystematics Research Institute, Ottawa, 
the Cornell University Museum, and the United States National Museum for loans of 
specimens including types. Particular thanks go to my friend and colleague Michael 
Shaffer at the British Museum for obtaining loans, dissecting type material, and offering 
his experience and advice at all stages of the work. 


LITERATURE CITED 


BYESZYNSKI, S. 1967. Studies on the Crambinae (Lepidoptera). Part 44. New neotropical 
genera and species. Preliminary checklist of neotropical Crambinae. Acta. Zool. 
Cracov. 12:39-110. 

BYESZYNSKI, S. & R. J. COLLINS. 1962. A short catalogue of the world species of the 

_ family Crambidae (Lepidoptera). Acta. Zool. Cracov. 7:197—389. 

Druce, H.H. 1896. Lepdioptera Heterocera: Geometridae, Pyralidae, and supplement 
to Sphingidae, pp. 273-336. In Godman, F. D. and O. Salvin (eds.), Biologia Centrali- 
Americana. Vol. II, pts. cxxvii-cxxxii. 

GASKIN, D. E. 1973. Revision of the New Zealand Chilonini (Lepidoptera: Pyralidae) 
and redescription of some Australian species. N. Z. J. Sci. 16:435-463. 

1985. Morphology and reclassification of the Australasian, Melanesian and 

Polynesian Glaucocharis (Lepidoptera: Crambinae: Diptychophorini). Aust. J. Zool. 

Suppl. Ser. 115:1-75. 

1986. Genus Diptychophora Zeller and a related new genus Steneromene from 
the neotropical region (Pyralidae: Crambinae). J. Lepid. Soc. 40:107-123. 

MEyYRICK, E. 1931. Exotic Microlepidoptera 4:33-192. 

1938. New species of New Zealand Lepidoptera. Trans. R. Soc. N. Zealand 67: 

426-429. 











288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OSTHELDER, L. 1941. Beitrag zur Kleinschmetterlings-Fauna Kretas. Mitt. Muench. 
Entomol. Ges. 31:365-370. 

ScHaAus, W. 1922. New species of Pyralidae of the subfamily Crambinae from tropical 
America. Proc. Entomol. Soc. Washington 24:127-145. 

WALKER, F. 1865. List of the specimens of lepidopterous insects in the collection of 
the British Museum, London 34:1097-1544. 

ZELLER, P. C. 1866. Beschreibung einiger amerikanischen Wickler und Crambiden. 
Entomol. Ztg. Stettin 27:137-157. 

1877. Exotische Microlepidopteren. Horae Soc. Entomol. Ross. 13:3-493, pls. 





1-6. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 288 


BOOK REVIEW 


BUTTERFLIES EAST OF THE GREAT PLAINS, by Paul A. Opler and George O. Krizek. 
1984. The Johns Hopkins University Press, 701 West 40th Street, Suite 278, Baltimore, 
Maryland 21211. Pp. 17 + 294. Price: $49.50 + shipping. 


The book gives comprehensive accounts of the butterflies of the thirty-one states east 
of the Great Plains. Detailed descriptions of more than 250 species are clearly and 
succinctly presented. The illustrative material, including detailed maps and 324 spectac- 
ular color photographs obtained in the field, add much to the species accounts. Some 
readers will no doubt complain that the 54 colored plates are set off in the central part 
of the book and not included with the species descriptions, but the very beauty of the 
photos as well as their taxonomic utility is best served by having them together. 

In the species accounts, the authors present the etymological derivation of the scientific 
name and a synopsis of the species, stating any noteworthy trait. A lengthier discussion 
and description of the butterfly follows, which includes its distinguishing characteristics, 
geographic variation, and various meaningful attributes including statistical measure- 
ments of both sexes. Both descriptive and map forms are used to present the overall 
range. Where applicable, they indicate temporary expansion of range beyond where the 
species is normally resident. A surprising number of species are indicated whose tem- 
porary northern extension of range is cut back by the severity of winter. The extensive 
review of county records contributed by more than one hundred lepidopterists helps 
make the data on distribution and habitat one of the most valuable contributions of the 
book. Habitat descriptions are provided for each species including specific vegetation, 
plant formations, and even associated soil types. 

The format for the species accounts also includes sections on life history and food 
sources, involving adult nectaring data as well as caterpillar host plants, and it is in these 
details that Opler and Krizek shine. The sections are full of data and challenging obser- 
vations which ought to provoke much more interest and enthusiasm among readers in 
adding to natural history observations. 

A lucidly written 33-page introduction contains a series of short essays on smaller 
topics reflecting the interests of field naturalists. Here the authors discuss such things as 
patterns of diversity, seasonality, and distribution. These sections should also whet the 
intellectual appetite and leave readers anxious to learn more. 

I find this book very revealing and exciting and feel it will be a valuable addition to 
the library of every lepidopterist, including those of “professional” as well as “amateur” 
standing. Part of its beauty lies in the obvious knowledge of field natural history possessed 
by the authors. It is well done! I hope they will favor us with another volume or two on 
the natural history of butterflies. 


JOHN C. DOWNEY, Graduate College, University of Northern Iowa, 138 Latham Hall, 
Cedar Falls, Iowa 50614. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 289-297 


BIOLOGY AND DESCRIPTION OF IMMATURE STAGES OF 
PHIGALIA STRIGATERIA (MINOT) (GEOMETRIDAE) 


LINDA BUTLER 


Division of Plant and Soil Sciences, P.O. Box 6108, 
West Virginia University, Morgantown, West Virginia 26506-6108 


ABSTRACT. Egg, larva, and pupa of Phigalia strigateria (Minot) are described for 
the first time from a population in eastern West Virginia. Eggs were deposited under 
loose bark of dead twigs. Thirty-two species of trees and shrubs were observed as larval 
hosts, with preference being shown for oaks, hickories, and common hackberry. Five 
larval instars and seven larval color forms were noted. Mean developmental time from 
egg to pupa was 28 days at 24°C. 


Four species of North American Phigalia belonging to the holarctic 
tribe Bistonini have been described (Rindge 1975). Three of the species, 
P. titea (Cramer), P. denticulata Hulst, and P. strigateria (Minot), are 
found only in eastern North America. Immature stages have been de- 
scribed only for P. titea (Butler 1985a, Talerico 1968). 

The distribution of P. strigateria was given by Rindge (1975) as 
eastern North America, southern Ontario, and Quebec, from the At- 
lantic Ocean to about longitude 100°W. While adult P. strigateria have 
been often recorded, almost no information is available on immature 
stages. Prentice (1963) reported a single larval collection from Ulmus 
americana L.. at Brockville, Ontario, in June 1950; he considered the 
species rare in that locality. 

Beginning in 1981, significant defoliation of hardwood forests in 
eastern West Virginia was attributed to a looper complex. Some larval 
collections were made in 1982, and a detailed study of the looper 
complex was conducted in 1983 and 1984. Phigalia strigateria was 
sufficiently abundant to provide material for biological and descriptive 
studies. Results of these studies are presented here. 


MATERIALS AND METHODS 


Study areas were in eastern West Virginia in an oak-hickory-pine 
forest on dry upland sites. Two study areas were on Cacapon Mountain 
(Cacapon State Park) in Morgan Co. One area, Batt Picnic Area (Batt), 
was at 381 m elevation, while the Cacapon Overlook area (Cac) was 
at 701 m. The Elkhorn Mountain (Elk) area was on the border of Grant 
and Hardy counties at 732 m elevation. During 1981-82, Batt was 
100% defoliated while the other sites were about 25% defoliated. De- 
foliation in 1983 was greatly reduced because populations collapsed, 
and by 1984 defoliation was sparse. 

Field studies were begun with observations and collections of emerg- 
ing adult P. strigateria on 17 March 1983. Samples of adults were 


290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


taken for species confirmation and collection of eggs. Oviposition habits 
were noted and fecundity was determined by allowing field-collected 
females to oviposit on dead twigs in the laboratory. All laboratory 
studies were conducted at 24°C and 12:12 photoperiod. Moths were 
not provided with food or water as the mouthparts are rudimentary. 

Larvae were collected in the field and their host plants recorded. 
Botanical nomenclature is that of Bailey and Bailey (1976). Stage du- 
rations were determined by rearing 60 larvae hatched from eggs laid 
in laboratory cages. Larvae were reared on leaves of sugar maple in 
groups of 10 in large Petri dishes. Larvae were checked daily and food 
was changed every other day. Descriptions were based on both labo- 
ratory-reared and field-collected larvae and pupae. Terminology fol- 
lows Hinton (1946) and McGuffin (1967). Larval head measurements 
were made with an ocular micrometer; early instars were measured at 
30x and later instars at 10x. Illustrations of larvae and pupae are by 
the author. Voucher specimens of larvae and adults are in the West 
Virginia University Collection. 


RESULTS 
Phenology, Life History, Food Plants 


In all study areas, the seasonal occurrence of P. strigateria was sim- 
ilar to that of P. titea (Butler 1985b). Adults were collected at Batt on 
17 March 1983 and at the higher altitudes of Cac and Elk beginning 
24 March. Males were most often observed resting on tree trunks, while 
females were climbing tree trunks, resting, or ovipositing on dead twigs. 
Female Phigalia taken at the study sites between 31 March and 26 
April were primarily P. titea. Females of P. strigateria made up the 
following percentage at each site: Batt 9% (21n), Cac 10% (29n), Elk 
8% (26n). 

Oviposition habits of P. strigateria are similar to those of P. titea; 
eggs are deposited most frequently under loose bark or in cracks or 
roughened areas of dead hardwood twigs. Females were observed ovi- 
positing in the field on dead twigs of Acer spp., Cornus florida L., 
Quercus spp., Betula lenta L. and Hamamelis virginiana L. Field- 
collected females (9n) lived two to four days in the laboratory and 
began ovipositing one to two days after being brought from the field. 
They oviposited a mean of 149 (range 26-319) eggs (9n). Females 
producing the fewest eggs had probably begun ovipositing before they 
were collected. 

Five larval instars were found for this species; mean developmental 
time from egg to pupa was approximately 28 days at 24°C (Table 1). 

Field larval development was similar to that of P. titea (Butler 1985a). 


VOLUME 40, NUMBER 4 291 


TABLE 1. Development time of P. strigateria reared on leaves of sugar maple at 
24°C. Means based on 60 larvae. 


Time in instar (days) 


Instar Mean Range 

i 4.1 3-5 

2 3.4 2-4 

3 3.4 2-5 

4 3.2 2-4 

) (er4 6-9 

Prepupa 6.5 5-9 
Total 27.8 21-34 


Egg hatch in 1983 began about 1 May at the lower altitude at Batt 
and about a week later at the higher altitudes. Larvae hung down on 
silk lines and ballooned away from dead twigs on which they hatched. 
Larvae fed for four to five weeks at each site before they moved into 
the soil to pupate. 

Larval populations at all three study sites in 1983 consisted largely 
of P. titea, ranging from 77 to 94% (2,162n) of the population at Cac 
and Elk, respectively. Larval percentages of P. strigateria at the study 
sites were: Batt, 4%; Cac, 8%; and Elk, 3%. Other species of larvae 
present in noticeable numbers were linden looper, Erannis tiliaria 
(Harris), 2-138%; and fall cankerworm, Alsophila pometaria (Harris), 
1-2%. 

Phigalia strigateria larvae were observed feeding on the following 
32 hosts: 


Juglans nigra L. Amelanchier canadensis (L.) 
Carya ovata (Mill.) A. x grandiflora Rehd. 

C. tomentosa Nutt. Malus sylvestris Mill. 

C. glabra (Mill.) M. coronaria (L.) Mill. 
Betula lenta L. Rubus spp. 

Corylus americana Marsh. Prunus serotina Ehrh. 
Quercus alba L. Cercis canadensis L. 

Q. prinus L. Acer negundo L. 

Q. stellata Wangenh. A. saccharum Marsh. 

Q. rubra L. A. rubrum L. 

Q. coccinea Muenchh. A. pensylvanicum L. 

Q. velutina Lam. Tilia americana L. 

Ulmus rubra Muhlenb. Parthenocissus quinquefolia (L.) Planch. 
Celtis occidentalis L. Nyssa sylvatica Marsh. 
Hamamelis virginiana L. Cornus florida L. 

Crataegus spp. Vaccinium augustifolium Ait. 


Host plants of P. strigateria are similar to those of P. titea (Butler 
1985b). 


292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Favored host plants appeared to be oaks, hickories, and hackberry. 
Only on hackberry was the percentage of P. strigateria larvae within 
the looper population markedly different from that previously men- 
tioned; P. strigateria larvae appeared to make up 40-70% of the looper 
population on hackberry observed at Batt and Elk. 

During 1983, high rates of parasitism by tachinids, ichneumonids, 
and braconids were observed in field-collected larvae of all four geo- 
metrid species. Parasitism was highest for P. strigateria, however, 
reaching 70-90% (135n) in some larval collections. While populations 
of all looper species in the study areas showed a dramatic collapse in 
1983, the collapse of P. strigateria was especially notable. In 1984, no 
adult P. strigateria were found, and larvae made up less than 0.01% 
(250n) of the population at each site. 


Description of Immature Stages 


Eggs. Eggs are sculptured, slightly rough textured, and oblong, with one end broadly 
rounded or blunt and the other end conical. They are yellow, becoming duller just before 
hatching. Empty P. strigateria chorions are a pale golden yellow in contrast to those of 
P. titea which are pale lavender. Eggs laid in the laboratory by field-collected females 
had a mean length of 0.75 mm (range 0.66-0.79) and mean width of 0.45 mm (range 
0.40-0.50) (80n). 

Larvae. Instars 1 through 3 are uniform in color. Larvae of the last two instars show 
much variability. The following descriptions are of coloration and mean head widths of 
instars of the most abundant color form of P. strigateria. 

Instar 1 (26n): Head (0.27 mm) pale yellow-brown; ground color of body pale greenish 
tan; pinnaculae black surrounded by white cuticle. Broad diffuse dorsal stripes and broad 
irregular lateral stripes are greenish white. Cervical shield pale brown; lateral shields of 
anal prolegs brown, prominent. 

Instar 2 (8n): Head (0.50 mm) pale yellow-brown with reddish brown granulations. 
Ground color of body dark greenish black. Pinnaculae not prominent; setae near spiracles 
each on a small white chalaza. Dorsal stripes greenish white, prominent when viewed 
with magnification, but very fine, irregular, and broken. Greenish white subdorsal and 
lateral stripes just above and below spiracle with diffuse grayish white fill around spiracle. 
Lateral stripe most prominent on abdominal segments 1-6. Ventral proleg greenish black. 
Anal plate grayish white with greenish black maculations; anal proleg shields prominent, 
color of ground, contrasting with paler color of anal segment. Mid-ventral pale stripe 
expanded at middle of each sternite. Secondary setae are present for the first time in this 
instar. 

Instar 3 (33n): Similar to instar 2. Head (0.81 mm) reddish brown. Dorsal and lateral 
stripes more prominent. Lateral stripe yellowish, extending down ventral proleg; absent 
beyond sixth abdominal segment. Anal plate and prolegs pale tan with brownish mac- 
ulations. 

Instar 4 (25n): Head capsule (1.36 mm) yellow, roughened with reddish brown mac- 
ulations; thoracic legs reddish brown; body ground color black. Dorsal stripes yellow, 
fine, irregular, often not continuous; black fill frequently between stripes but not consis- 
tent; stripes most prominent on abdominal segments 1-2 and 6-9. Dorsal chalazae of 
eighth abdominal segment most prominent. Lateral stripe irregular, yellow, generally 
broad and most prominent on abdominal segments 1-6, weak on thorax; on abdominal 
segment 6, stripe extends down proleg and terminates; lateral striping on posterior ab- 
dominal segments appearing only as yellow flecks. Spiracles with black peritreme; spi- 
racular valve yellow. Cervical shield black with brownish tan irregular fill. Anal and 


VOLUME 40, NUMBER 4 293 


TABLE 2. Percentage of color forms of 4th and 5th instar P. strigateria collected 
between 17 May and 8 June 1983. 


Percentages 
No. Yellow/ 

Study site larvae Typical Dark Pale stripe Yellow Orange Brown 
Batt 97 69 1 16 aI: 2, 0 1 
Cac 139 35 9) 19 26 i 6 2 
Elk 60 40 0 19 18 5 18 0 


proleg shields prominently mottled yellow-brown. Venter yellowish brown with pale 
yellow mid-ventral stripe. 

Instar 5 (34n): Head capsule (2.24 mm) as in instar 4. Legs tan and dark brown. Dorsal 
stripes well developed on first nine abdominal segments, partially developed on meta- 
thorax, as dashes only on mesothorax, absent on prothorax; stripes yellow, often giving a 
chain-link appearance dorsally. Cervical shield and anal prolegs granulate dull brown; 
anal plate pale tan with small, dark maculations. Venter with a broad tan stripe expanded 
on each segment; stripe on abdominal sternites 7-8 often paler. Other features similar 
to those of 4th instar. 

The most frequently observed variations in color patterns of instars 4 and 5 were the 
pale form and yellow form with subdorsal stripe. 

Pale form—Yellow striping more intense. Dorsal stripes broad, pale yellow with mid- 
dorsal dark dashes only on thorax; only a narrow band of black above the broad yellow 
lateral stripe. Dashed black subventral line; legs brown with dark coxae. Cervical and 
anal shields yellow with brown maculations. 

Yellow form with subdorsal stripe—Head paler than on darker forms. Ground color 
of body yellow. Dark points include only a prominent black subdorsal stripe which is 
continuous from prothorax through abdominal segment 10 or continuous only to abdom- 
inal segment 5, and chalazae on abdominal segments 2, 8, and 8, which are often black. 

Other variations included a yellow form without subdorsal stripes; an orange form, 
similar to the typical form but with orange replacing yellow; a dark form with all pale 
patterns reduced; and a brown form. 

Samples of 4th and 5th instar P. strigateria collected at the study sites from 17 May 
through 8 June consisted of the forms in Table 2. 

Chaetotaxy of last instar: Head with P1 somewhat in front of P2; A2 in front of and 
below A8 and above Al; L1 directly above 02; 01 directly posterior to ocellus 3; AF1 
and AF2 widely separated; F1 almost directly below AF1 and above and in front of C2; 
Cl close to C2 and on edge of adfrons; labrum with L2 and M2 longer than other labral 
setae; 02 and Cl longest of all head setae (Figs. 1, 2). Prothorax with XD1, XD2, D1, 
D2 and SD2 approximately equidistant from each other; SD1 very small and close to 
SD2; L2 small and directly below L1: SV1 longer than SV2 (Fig. 8). On mesothorax D1, 
SD2 and L3 in a vertical line; D2 and SD1 anterior; SD1 very small; L2 smaller and 
somewhat anterior to L1; SV1 relatively long (Fig. 8). On the second abdominal segment, 
D2 is slightly above D1; SD1, L1 and L2 equidistant from spiracle; SD1 and L2 in front 
of spiracle; L1 behind spiracle; L3 below and slightly posterior to L2; SV3 behind and 
level with L8; SV1 below and in front of SV4 (Fig. 7). Sixth abdominal segment with 
D1 above D2; SD1 directly below D1 and above anterior margin of spiracle; L2 directly 
below SD1 and just below spiracle; L1 behind spiracle and slightly more removed from 
spiracle than are SD1 and L2; L3 in front of L2 (Fig. 6). On eighth abdominal segment, 
D1 on prominent tubercle; SD1 and D2 level with each other; SD1, L2 and L1 arranged 
around spiracle as on sixth abdominal segment; SV3 slightly anterior to L1; SV1 directly 
below SV3 (Fig. 6). Abdominal segment 9 with D1, SD1 and L1 each somewhat anterior 
to the seta above; SV1 posterior to L1 (Fig. 6). Segment 10 with SD1 and D1 widely 
separated; D2 equidistant between L1 and adjacent D2; CP1 above CP2; Lgl, Lg2 and 
Lg3 in vertical line; CD1 above CD2 (Figs. 6, 11). On abdominal segments 1 and 2, 1 


294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





ns 


Fics. 1-5. Phigalia strigateria larval head structures. Scale line length in parentheses. 
1, Lateral view of head (0.5 mm); 2, Frontal view of head (0.5 mm); 3, Ventral view 
of mentum, hypopharynx, labial palpi, spinneret and maxilla (0.25 mm); 4, Inner view 
of right mandible (0.25 mm); 5, Ventral view of maxilla (0.25 mm). 


ventral seta is present; on segment 8, SV1 and SV4 closely adjacent (Fig. 12). The 
following chalazae are most prominent on last instar larvae: D2 and L1 on abdominal 
segments 2 and 3, and D1 on segment 8. Secondary setae numerous, especially dorsally 
and laterally on all segments; setae are fine, irregular and about % the length of primary 
setae. Body roughened with microspines anteroventrally on prothorax and ventrally on 
mesothorax. 


VOLUME 40, NUMBER 4 295 





Fics. 6-8. Phigalia strigateria larva. Scale line = 0.5 mm. 6, Lateral view of abdom- 
inal segments 6-10; 7, Lateral view of abdominal segment 2; 8, Lateral views of pro- 
and mesothorax. 


Mouthparts: Mandibles with 4 large and 5 small teeth, more basal mandibular seta 
longest (Fig. 4); postmentum with a pair of long setae (Fig. 3); hypopharynx heavily 
sclerotized; spinneret tubular in shape, slightly tapering apically; labial palps almost the 
length of spinneret (Fig. 3); ventral side of each maxilla with 4 prominent setae, most 
apical one smallest (Fig. 3); terminal lobe of maxilla with 3 setae and 2 elongated papillae, 
apicalmost seta longest (Fig. 5). 

General: Last instar about 26 mm long and 3.1 mm wide; thoracic leg claw dark 
brown, pointed with 1 dorsal simple seta and 3 lateral and ventral bladelike setae (Fig. 
18); A6 crochets a biordinal mesoseries in unbroken band. 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





ioe 
SS 
— 13 


Fics. 9-13. Phigalia strigateria larva and pupa. Scale line length in parenthesis. 9, 
Pupa, ventral view (0.5 mm); 10, Pupal cremaster (0.25 mm); 11, Anal plate (0.5 mm); 
12, Ventral view of Al and A2 (0.5 mm); 13, thoracic leg claw (0.1 m). 


Pupa. Reddish brown; eyes large, completely exposed, rounded; labrum hexagonal; 
maxillae slightly shorter than antennae; prothoracic leg extending about % length of 
maxilla, prothoracic femur not exposed; mesothoracic leg ending near antennae; meta- 
thoracic legs exposed beyond apex of maxillae (Fig. 9). Cremaster always bifurcate, spines 
usually asymmetrical (Fig. 10). Segment 7 constricted apically; abdominal segments 
coarsely punctate. Length similar for male and female, mean 9.9 mm (range 6.5-12) 
(43n); female stouter. 


DISCUSSION 


The chaetotaxy of mature P. strigateria larvae shows some variation 
in location of the following setae: P1 and A2 setae on head more medial 
than shown; L2 on abdominal segment 2 slightly before spiracle; anal 
plate with D2 setae closer to each other than to their respective L1’s. 


VOLUME 40, NUMBER 4 297 


Comparison of P. strigateria larvae with those of P. titea from the 
same study areas in West Virginia showed close similarity in location 
of primary setae between the two species. Primary setae of P. titea are 
longer with most arising from small but well defined chalazae. SD1 
and L2 on the prothorax are smaller in P. titea than in P. strigateria. 
Secondary setae of P. titea are shorter and sparser than those of P. 
strigateria. All body cuticle of P. titea is densely covered with mi- 
crospines. Because of the marked color differences in larvae of the two 
species, they cannot be confused. 

The pupa of P. titea is larger than that of P. strigateria (Butler 
1985a). The labral shapes are more rounded than those of P. titea. 
Phigalia titea vertex lacks the prominent rugosity of P. strigateria; 
frontoclypeal area is smooth in P. strigateria, rugose in P. titea. 


ACKNOWLEDGMENTS 


I thank Superintendent Philip Dawson and his staff of Cacapon State Park for their 
cooperation during this study; Mr. and Mrs. Fred Riggleman of Dorcas, West Virginia, 
for allowing me to conduct a portion of this research on their land at Elkhorn Mountain; 
Vicki Kondo, Terry Stasny, and Beth Cahape for laboratory assistance; and James W. 
Amrine, John E. Hall, and Joseph E. Weaver for comments on the manuscript. 

This paper is published with the approval of the West Virginia Agricultural and 
Forestry Experiment Station as Scientific Article No. 1941. 


LITERATURE CITED 


BAILEY, L. H. & E. Z. BatLey. 1976. Hortus third, a concise dictionary of plants 
cultivated in the United States and Canada. Macmillan, New York. 1290 pp. 

BUTLER, L. 1985a. Biology of the half-wing geometer, Phigalia titea Cramer (Geo- 
metridae), as a member of a looper complex in West Virginia. J. Lepidop. Soc. 39: 
177-186. 

1985b. Food plant studies for the half-wing geometer, Phigalia titea Cramer 
(Lepidoptera: Geometridae). Can. Entomol. 117:547-551. 

HINTON, H. W. 1946. On the homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Royal Entomol. 
Soc. 97:1-37. 

McGuFFIn, W. C. 1967. Immature stages of some Lepidoptera of Durango, Mexico. 
Can. Entomol. 99:1215-1229. 

PRENTICE, R. M. 1963. Forest Lepidoptera of Canada. Publ. Canada Dept. Forestry, 
Forest Entomol. Pathol. Br., No. 1018, Vol. 3, pp. 263-543. 

RINDGE, F. H. 1975. A revision of the new world Bistonini (Lepidoptera: Geometridae). 
Bull. Am. Mus. Nat. Hist. 156:71-155. 

TALERICO, R. L. 1968. Life history of the looper Phigalia titea in Virginia. Ann. 
Entomol. Soc. Am. 61:557-561. 





Journal of the Lepidopterists’ Society 
40(4), 1986, 298-303 


TRAP PREFERENCES OF RETINIA METALLICA AND 
SEASONAL FLIGHT BEHAVIOR OF RETINIA SPP., 
RHYACIONIA SPP. (TORTRICIDAE), AND 
CHIONODES SPP. (GELECHIIDAE) 

IN THE DAKOTAS 


MARY ELLEN DIX 


Rocky Mountain Forest and Range Experiment Station, 
Forestry Sciences Laboratory, East Campus, University of Nebraska, 
Lincoln, Nebraska 68583 


AND 


MARTIN JACOBSON 


Biologically Active Natural Products Laboratory, 
Agricultural Environmental Quality Institute, Agricultural Research Service, 
Beltsville, Maryland 20705 


ABSTRACT. At high population levels, white and green traps baited with (Z)-7- 
dodeceny] acetate caught more Retinia metallica than blue traps. Diamond-shaped traps 
were more effective than cup traps, but did not differ significantly in effectiveness from 
triangular-shaped traps. Rhyacionia fumosana and R. neomexicana responded through- 
out May to synthetically-baited traps. Rhyacionia bushnelli and Retinia metallica flew 
in late May or early June. 


Tip mining lepidopterous larvae can cause extensive damage to pon- 
derosa pine (Pinus ponderosa Laws.) in the northern Great Plains (Stein 
& Kennedy 1972). Information on their distribution is limited; effective 
techniques for detecting and evaluating infestations are not available 
(Dix et al. 1984). Accurate determination of the flight period of Lep- 
idoptera that are concealed for most of their life cycle is crucial to the 
effective timing of insecticide applications. Sex attractants are ideal for 
detecting and delineating the adult flight period of such Lepidoptera 
(Stevens et al. 1980). Jacobson and Jennings (1978) and Stevens et al. 
(1980) identified attractants of Rhyacionia fumosana Powell and Rhy- 
acionia neomexicana (Dyer). Several possible lures of Retinia metal- 
lica (Busck) and Rhyacionia bushnelli (Busck), two of the more com- 
mon species in the north central U.S., were identified by Dix et al. 
(1984). A trapping technique for Retinia metallica using these lures 
needs to be refined. This article describes trap design and trap color 
preferences of R. metallica, and delineates the flight period of Retinia 
spp., Rhyacionia spp. (Tortricidae: Olethreutinae), and Chionodes spp. 
(Gelechiidae) that infest ponderosa pine in North Dakota and South 
Dakota. 


VOLUME 40, NUMBER 4 299 


TABLE 1. Description of sites used to determine flight periods of pine-feeding Lep- 
idoptera in North Dakota and South Dakota, 1973-76. 


Esti- 


e : mated ; 
Year Locality tari eee ane Seog : 
1975 Near Burning Coal Vein, Little Native 05-20 500 10 May- 
Missouri Grasslands, Custer Na- 20 July 
tional Forest, Slope Co., North 
Dakota 
North Cave Hills, Custer National Native 0.5-20 500 10 May- 
Forest, Harding Co., South Da- 20 July 
kota 
Slim Buttes, Custer National For- Native 0.5-20 500 10 May- 
est, Harding Co., South Dakota 20 July 
Big Sioux Conifer Nursery, Cod- Planted 2-6 3,000 8 May- 
dington, Co., South Dakota 15 June 
1976 Near Burning Coal Vein, Little Native 0.5-20 500 10 May-— 
Missouri Grasslands, Custer Na- 3 Aug 
tional Forest, Slope Co., South 
Dakota 
Big Sioux Conifer Nursery, Cod- Planted 2=6 8,000 27 May-— 
dington Co., South Dakota 22 June 


METHODS AND MATERIALS 


Trap design and trap color preferences of R. metallica. Trials were 
conducted in May and June at the Big Sioux Conifer Nursery, Cod- 
dington Co., South Dakota. Initially, a moderate population (5 pitch 
blisters/tree) was present in the 9-year-old ponderosa pine provenance 
planting, and a high population (36 pitch blisters/tree) was present in 
the 20-year-old ponderosa pine in the nursery’s border planting. 

A cardboard cup trap (0.24 liter), a triangular milk carton trap (9 
em high and 15 cm long), and a diamond-shaped milk carton trap (9 x 
9 x 15 cm) were each lined with Stikem Special® and were baited 
with 10 mg of (Z)-7-dodecenyl] acetate. All traps were open at both 
ends and were white. There were six blocks of three traps (one of each 
design). Traps in each block were hung in the border planting 20 m 
apart at a height of 1.5 m. 

The effect of trap color on trap catch was determined with diamond- 
shaped traps painted white (2A1), pale blue (24A5), green (28D8), or 
fluorescent orange (7A8). Notations following colors refer to Kornerup 
and Wanscher’s (1967) standard colors. All traps were baited with 
rubber septum dispensers containing 10 mg (Z)-7-dodecenyl acetate 
plus 10 mg trioctanoin and were deployed 20 m apart at a height of 
1.5 m in perimeter ponderosa pines. There were six blocks of four traps 
(one of each color) in trial 1 and four blocks of four traps in trial 2. 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Comparison of trap catch of male R. metallica with different sticky trap 
designs baited with 10 mg of (Z)-7-dodeceny] acetate plus 10 mg trioctanoin. 


No. males/trap No. males/cm? 
Surface area Number = ee ee 
Design (cm?) of traps Mean! SE Mean! SE 
Diamond 427.5 3 187a 21.4 0.5a 0.05 
Triangular 405.0 3 142ab 28.4 0.3a 0.06 
Cup 220.5 3 59b 4.3 0.3a 0.02 


1 Means followed by the same letter are not significantly different (P < 0.05) according to Tukey’s procedure for 
multiple comparisons. 


Each week for three weeks, trap catches were counted and the traps 
were randomly reassigned to a new site within the block. Analysis of 
variance and Bartlett’s test for homogeneous variance were performed 
on the total catch per trap. When necessary, the data were transformed 
by In(x + 1) to stabilize the variance for analysis of variance. Tukey’s 
honestly significant difference procedure at the 5% level was used to 
separate means (Sokal & Rohlf 1969). 

Seasonal flight record. During spring and early summer of 1975 
and 1976, known attractants of Rhyacionia spp. and Chionodes spp.., 
and several related synthetic compounds were used to attract Rhy- 
acionia spp., Retinia spp., and Chionodes spp. that damaged ponderosa 
pine in native and planted stands (Table 1). Compounds used as lures 
included (E)-7-decenyl acetate, (Z)-7-deceny] acetate, (E)-7-dodeceny] 
acetate, (Z)-7-dodeceny! acetate, (Z)-7-dodecen-1-ol, (E)-8-dodeceny] 
acetate, (Z)-8-dodecenyl acetate, (Z)-9-dodeceny]l acetate, (E)-9- 
dodeceny] acetate, (E)-9-dodecen-1-ol, (Z)-10-dodecenyl acetate and 
(E)-10-dodenceny] acetate. In 1975, each compound was replicated 
twice at a site, and traps and dispensers were changed three times 
during a trapping period. In 1976, each compound was replicated five 
times at a site, and traps were changed six times at two-week intervals 
during a trapping period. 

Traps were cardboard cups (0.24 liter) with a 2.5 cm diameter open- 
ing at both ends. The inside was coated with Stikem Special®. In 1975, 
a cotton wick dispenser (12 x 10 mm) was impregnated with 20 mg 
of a test compound and placed in the bottom of each trap. In 1976, a 
rubber septum dispenser (5 x 9 mm) baited with 10 mg of test com- 
pound was used. In 1975 and 1976, compound activity was prolonged 
by adding 10 mg of the extender trioctanoin. Control dispensers also 
were baited with 10 mg of trioctanoin and were replicated twice per 
site in 1975 and five times per site in 1976. Traps baited with synthetic 
attractants or with control dispensers were hung on ponderosa pine 
branches at a height of 1.5 m and a spacing of at least 20 m. 


VOLUME 40, NUMBER 4 301 


TABLE 8. Effect of trap color on catches of male Retinia metallica in diamond-shaped 
sticky traps baited with 10 mg of (Z)-7-dodeceny] acetate plus 10 mg trioctanoin. 


Trial 1 Trial 2 
te No. males/trap Hee ae No. males/trap 
Trap color of traps Mean! SE of traps Mean! SE 
White 4 150.3a 39.91 4 3.5a 1.19 
Green 4 123.5a 34.50 4 2.5a 0.95 
Orange 4 95.3a 44.18 + 1.8a 0.63 
Blue 4 16.0b 2.16 4 0.8a 0.48 


1 Means followed by the same letter are not significantly different (P < 0.05) according to Tukey’s procedure for 
multiple comparisons. 


Captured moths were identified and counted when traps were 
changed. Moths were removed from traps with forceps or by carefully 
cutting the trap around the moth. Specimens were tentatively identi- 
fied and numbers of moths per species were recorded. Representative 
samples of each presumed species were sent to specialists for identifi- 
cation. 


RESULTS AND DISCUSSION 


Trap design and trap color preferences of R. metallica. Traps in 
several blocks were blown down during a spring storm; catches in traps 
from these blocks were not included in analyses. The second color 
preference trial was conducted the following spring, after the number 
of active R. metallica had abruptly decreased (less than 1 pitch blister / 
tree). An unusually wet August and winter probably contributed to the 
high larval mortality and reduction in number of adults. 

Effectiveness of the traps varied with design and color (Tables 2, 3). 
Diamond-shaped traps had the largest surface area and caught more 
moths than either triangular or cup traps. However, only mean catches 
per trap of the diamond-shaped traps and cup traps differed signifi- 
cantly. At high population levels, as exhibited in trial 1, white, green, 
and orange traps caught significantly more males than blue traps (Ta- 
ble 3). However, at low population levels (trial 2), trap color did not 
affect trap catch. 

Seasonal flight behavior. The beginning and end of male flight for 
five species of Olethreutinae which infest the branch tips of ponderosa 
pine varied yearly with the onset of spring. For example, the spring 
of 1976 was warmer than in 1975, and moth flights of Retinia metal- 
lica, R. gemistrigulana and Rhyacionia fumosana were earlier (Fig. 1). 

In southwestern North Dakota and northwestern South Dakota, male 
R. fumosana and R. neomexicana, the first species trapped in the 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1 1975 Slope Co., N. Dak. 

gm (975 North Cave Hill, Harding Co., S. Dak. 
1975 Slim Buttes, Harding Co., S. Dak. 
E33 1975 Coddington Co., S. Dak. 

CJ 1976 Slope Co., N.Dak. 

ZZ! 1976 Coddington Go., S.Dak. 


Retina metallica a 
ee | 


Retina gemistrigulana 


Rhyacionia bushnelli 
Rhyaconia fumosana 
Rhyacionia neomexicana 


Chionodes spp. 





Fic. 1. Seasonal flight of Tortricidae: Olethreutinae and Gelechiidae that damage 
ponderosa pine in North Dakota and South Dakota. 


spring, were usually caught in early May. Rhyacionia bushnelli, Re- 
tinia metallica, and R. gemistrigulana (Kearfott) were caught two to 
three weeks later, frequently in late May or early June. Large numbers 
of Chionodes spp. (Gelechiidae), which mine cones of ponderosa pine, 
were caught during June and July. Flight of Retinia metallica in north- 
eastern South Dakota (Coddington Co.) was about two weeks earlier 
than in northwestern South Dakota. 

In conclusion, white, green, or orange diamond- and triangular- 
shaped traps are effective in catching male R. metallica. However, 
diamond-shaped traps are preferred for future attractant trials and for 
detecting males because they provide a larger surface area. Time of 
moth flight varies annually. Additional research is needed to determine 
the effect of cumulative atmospheric and ground temperature on moth 
flight and to develop a method for accurately predicting moth flight. 


ACKNOWLEDGMENTS 


William Miller, USDA Forest Service, North Central Experiment Station, St. Paul, 
Minnesota, and Jerry A. Powell, University of California, Berkeley, identified the Tor- 


VOLUME 40, NUMBER 4 303 


tricidae; and R. W. Hodges, USDA, Agricultural Research Service, Northeastern Regional 
Agricultural Research Center, Beltsville, Maryland, identified the Gelechiidae. 


LITERATURE CITED 


Dix, M. E., A. D. TAGESTAD, J. D. STEIN & M. JACOBSON. 1984. Detecting tip mining 
Olethreutinae (Tortricidae) moths in the northern and central Great Plains with 
synthetic attractants. USDA Forest Service Research Note RM-445. Rocky Mountain 
Forest and Range Experiment Station, Fort Collins, Colorado. 5 pp. 

JACOBSON, M. & D. T. JENNINGS. 1978. Attraction of Rhyacionia neomexicana (Dyar) 
to synthetic pheromones. J. Environ. Sci. Health, Part A, 13:429-443. 

KORNERUP, A. & J. H. WANSCHER. 1967. Methuen handbook of colour. 2nd ed. Meth- 
uen and Co. Ltd., London. 243 pp. 

SOKAL, R. R. & F. J. ROHLF. 1969. Biometry: The principles and practice of statistics 
in biological research. W. H. Freeman and Co., San Francisco. 776 pp. 

STEIN, J. D. & P. C. KENNEDY. 1972. Key to shelterbelt insects in the northern Great 
Plains. USDA Forest Service Research Paper RM-85. Rocky Mountain Forest and 
Range Experiment Station, Fort Collins, Colorado. 153 pp. 

STEVENS, R. E., C. SARTWELL, JR., T. W. KOERBER, G. DATERMAN, L. L. SOWER & J. A. 
POWELL. 1980. Western Rhyacionia (Lepidoptera: Tortricidae, Olethreutinae) pine 
tip moths trapped using synthetic sex attractants. Can. Entomol. 112:591-603. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 304-314 


LIFE HISTORY OF NEMORIA GLAUCOMARGINARIA 
(BARNES & MCDUNNOUGH) AND LARVAL 
TAXONOMY OF THE TRIBE NEMORIINI 
(GEOMETRIDAE: GEOMETRINAE) 


ADAM H. PORTER 


Department of Zoology, University of California, 
Davis, California 95616 


ABSTRACT. Morphology, chaetotaxy, and color pattern of immature Nemoria glau- 
comarginaria (Barnes & McDunnough) are described for the first time. Large dorsolateral 
projections on the middle abdominal segments provide crypsis on the host plant (Quercus 
spp.), enhanced by a “shaking” behavior while moving. Larval color pattern of N. dar- 
winiata punctularia Barnes & McDunnough is also figured, and two new hosts are 
reported (Arctostaphylos sp. and Ceanothus cordulatus Kell.). The description of glau- 
comarginaria is used as the basis for recognition of the genus Nemoria. A preliminary 
key to larvae of the four genera in the tribe Nemoriini is presented, based on fourteen 
species. 


Taxonomy of the Geometrinae based on adult genital morphology 
has been covered in detail by Ferguson (1969, 1985). His studies reveal 
a remarkable array of parallelisms and convergences in the facies of 
adults. Individuals from different species groups within genera, or even 
from different tribes, have routinely been classified as conspecific in 
major museum collections. For example, Nemoria glaucomarginaria 
(Barnes & McDunnough) of the bistriaria group is often found with 
N. darwiniata (Dyar) of the obliqua group, and species of Dichor- 
dophora Prout (Dicherdophorini) are often found with Dichorda War- 
ren (Nemoriini). Although the taxonomic groupings outlined by Fer- 
guson are well defined, their interrelations remain unknown. Adult 
characters present numerous problems for phylogenetic reconstruction: 
the “superficial” characters show many parallelisms, convergences, and 
plesiomorphies, while the genitalia are so different between tribes and 
so similar within genera that most phylogenetic information is lost. 
However, the larvae show remarkable adaptations for crypsis, and may 
give information of phylogenetic significance not present in the adults. 

This paper describes the larva and life history of Nemoria glauco- 
marginaria and uses this information as the basis for the generic rec- 
ognition of Nemoria larvae. It concludes with a preliminary key to the 
genera of nemoriine larvae. 


MATERIALS AND METHODS 


Gravid females of N. glaucomarginaria and N. darwiniata punc- 
tularia Barnes & McDunnough were collected at ultraviolet and visible 
wavelength fluorescent lights from the following California localities 
in 1984 and 1985: glaucomarginaria: Gold Run, Placer Co.; Washing- 


VOLUME 40, NUMBER 4 305 


ton, Nevada Co.; Rubicon River Cyn., 1.5 km N Uncle Tom’s Cabin, 
El Dorado Co.; Barton Flats, San Bernardino Nat. For., San Bernardino - 
Co.; d. punctularia: Santa Rosa, Sonoma Co.; Baxter, Placer Co.; Meeks 
Bay, El] Dorado Co.; Carson River Cyn., Alpine Co.; Mill Ck. (Lundy 
Cyn.), Mono Co. Additional preserved material of d. punctularia was 
examined from Ash Cyn., Huachuca Mts., Cochise Co., Arizona. 

Ova were collected from moths confined in inflated, resealable plas- 
tic sandwich bags. Larvae were reared on foodplant cuttings in similar 
containers at 25°C under natural photoperiods throughout spring and 
summer. Specimens of each stage and instar were preserved in KAAD 
before transfer to 95% ethyl alcohol. Color photographs were taken of 
the ultimate instars. Figures were drawn with the aid of a camera 
lucida, with color patterns added freehand from the color slides. Chae- 
totaxy follows McGuffin (1964, 1967, 1977). 

More than 100 larvae were reared to adults. Five specimens of each 
stage were examined for morphological details. Colors are described 
from two specimens of each species exhibiting the most common col- 
oration. Colors should not be relied on for specific identification. 

Larvae of additional species used in constructing the key are from 
personal rearings or from the collection of Noel McFarland. The species 
examined were Nemoria pulcherrima (Barnes & McDunnough), N. 
unitaria (Packard), N. arizonaria (Grote), N. festaria (Hulst), Chlo- 
rosea margaretaria Sperry, C. roseitacta Prout, C. banksaria gracearia 
Sperry, Dichorda consequaria (Hy. Edwards), D. illustraria (Hulst), 
and Dichordophora phoenix (Prout). An additional undetermined lar- 
va from near Valyermo, Los Angeles Co., California, was also exam- 
ined; it is probably N. intensaria (Pearsall). Specimens used in for- 
mulating these descriptions will be deposited in the Bohart Museum 
(University of California, Davis) and in the McFarland Collection. 


Nemoria glaucomarginaria (Barnes & McDunnough) 


Egg. Ovate, 0.7 mm long by 0.5 mm wide, with flattened, slightly concave top; vertical 
sides 0.2-0.3 mm thick, with sharp edge between sides and top; decorated with small 
ridges in an irregular reticulate hexagonal pattern. Yellow when laid, gradually dark- 
ening through orange to dark red over a period of 9-12 days, becoming translucent 2 
days before hatch. Larva emerges through hole chewed in side wall at anterior end. A 
photograph of the related N. darwiniata punctularia (Comstock & Henne [1940], as 
pistacearia [Packard]) is representative of glaucomarginaria, as well as all Nemoria 
examined to date. Details of microstructure of glaucomarginaria eggs were not exam- 
ined, but seem to agree with those of N. bistriaria rubromarginaria (Packard) (Salkeld 
1983:153). Clutches of up to 55 eggs were recorded. 

First instar. Similar to ultimate instar (Fig. 1), but with dorsolateral projections on 
A2-4 smaller, thicker, and squared off; setae L1 and L2 at tips. Projections half as wide 
as segments anteroposteriorly (Dichorda are % as wide in the first instar). Color mottled 
brown, pattern obscure; setae clear and blunt. Head width 0.3 mm, body length 3 mm 
at hatch. Chaetotaxy not examined. 

Second instar. Like first instar, except dorsolateral projections on A2-4 slightly more 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





5 


Fics. 1-5. Nemoria glaucomarginaria, scale 1 mm, except as indicated. 1, Mor- 
phology of fifth instar, lateral view; 2, Head of fifth instar. Pattern of larger spicules and 
darker markings at right, chaetotaxy at left. Setae translucent; 3, Right mandible. Inter- 
nal aspect unshaded, external shaded. Scale 0.5 mm; 4, Anal shield, dorsal view; 5, A3, 
dorsal view, showing placement of setae and spiracle. 


elongate at L2 than Ll. Color mottled brown, gross elements of fifth instar pattern 
sometimes visible posteriorly and dorsally. 

Third instar. Structurally identical to fifth instar; coloration as in second instar, except 
pattern elements usually more visible. 

Fourth instar. Identical to fifth instar, except in size. 

Fifth instar. The following description is given by segment. 

Head (Fig. 2) width 2.1-2.5 mm; tan to brown, mottled with darker pigment as shown; 
covered with spiculiferous projections containing white pigment spots under a clear 
cuticle, spicules reaching greatest devlopment on dorsal epicranium. Setae translucent, 
not black as shown. Clypeus lighter than head, with wrinkles as in d. darwiniata (Dyar 
1904). Mandibles as illustrated (Fig. 3). 

The following descriptions refer to Figs. 1, 8, and 10, except as indicated: 

T1 with dorsal conical projections bearing setae XD1 and XD2; caudad another pro- 
jection bearing setae D1 and D2 together. Another swelling dorsolaterally bears setae L1 
and L2; SV2 and SV3 on swelling above leg. L2 long and hairlike, probably acting as a 
proprioceptor. Pattern as figured, colors light to very dark brown, with whitish markings; 
addorsal line often dark green. T2 with large protuberance elevating setae L1-3, SD1, 
and SV3; SV2 on swelling above leg. SD2 long and thin. Color as in Tl. T3 structure as 


VOLUME 40, NUMBER 4 307 





Fics. 6, 7. Nemoria glaucomarginaria pupa. 6, Ventral aspect; 7, Lateral aspect. 


in T2; green, resembling color of foodplant, especially posteriorly, with lighter markings 
as shown; protuberance light brown, dorsal markings green or brown. 

Al with D1 barely raised, closer to mid-dorsal line than D2. Lateral protuberance 
small, bearing SD2 only. Spiracle below protuberance, above L2 and L1. Color green 
with markings as shown; darker green shading below protuberance. Anterior mid-dorsal 
patch dark. A2-4 (Fig. 5) with D1 as in Al, except raised more. Spiracle above projec- 
tion; MD1 thin, anterior to spiracle, probably a proprioceptor. SD setae absent. Dorso- 
lateral projections large, conical, slightly bidentate, bearing L2 at greatest extremity, L1 
caudad; projections longer and more pointed on A8. SV4 on line between L3 and SV2 
in segments A3 and A4. Pattern as shown; color brown on anterior part of segments and 
on distal half of projections, green elsewhere. A2 often completely brown as shown, A8 
usually mostly green. White markings around anterior mid-dorsal patch, on caudal edge 
of projections, anterior subdorsal patch, and lateral line, pronounced in intersegmental 
areas; smaller whitish markings as figured. Adventral line dark, chainlike, often obscure. 
A5 with D1 placed further from mid-dorsal line than on A1-—4, still closer to mid-line 
than D2. SD setae absent. Lateral protuberance small, raising L1 and L2. Spiracle above 
protuberance. Color green, pattern as on A2—4, except anterior mid-dorsal patch obscure. 
A6 with D1 and D2 equidistant from mid-dorsal line. SD setae absent. Small swelling 
below spiracle bearing L1 and L2. Crochets reduced to half length in center, not wholly 
interrupted. Pattern as shown; anterior green, posterior brown to purplish brown with 
frosty countershading on leg. A7 with D1 and D2 equidistant from mid-dorsal line. SD 
setae absent. Small swelling below spiracle elevating L1 and L2. Pattern as shown; color 
light brown on swelling to dark brown generally, markings white; brown countershades 


308 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 8. Nemoria glaucomarginaria. Lateral aspect showing pattern of markings on 
fifth instar. Refer to text for color description. Scale 1 mm. 


to whitish ventrally. A8 with D1 raised dorsally on high conical projection directed 
caudally. Pattern as shown; projection light brown with whitish tip, dark chestnut brown 
on patch below this. Purplish brown below white subdorsal line, fading to whitish ven- 
trally. A9 with D1 on large chalaza; other setae as shown. A10 with anal plate as shown 
(Fig. 4). Sclerotized shield of proleg marked with darkly pigmented indentations of 
various sizes (Figs. 1, 10), slightly variable in placement. An additional seta, herein named 
LG4, present on the anteroventral corner of shield (also present in all other Nemoria 
examined). Pattern as shown; markings white to buff on brown background. Crochets 
wholly interrupted in center by fleshy pad. 

Entire surface of larva covered with spicules, each usually with white spot of pigment 
inside. They reach greatest size dorsally and on projections and protuberances, approach- 
ing length of blunt setae. Spicules smaller below, giving rugose, granular or velvety 
appearance; most without white pigment on segments A6-10. Most setae (except MD1) 
raised on chalazae (Fig. 10), which often accumulate spicules, especially dorsally. 

A loose cocoon is spun on the foodplant along a twig, incorporating pieces cut from 
nearby leaves, or from cage material (such as cloth or paper). 





g —\, 


Fic. 9. Nemoria darwiniata punctularia. Lateral aspect showing pattern of markings 
on fifth instar. Refer to text for color description. Scale 1 mm. 


VOLUME 40, NUMBER 4 309 


y) 
=e L1 S91 
L1 eae 
L2 
SV3 : v3 


y Isv2 
mvi2 
> MV2 sv4 


Pa 5 


T2 Na 








Fic. 10. Setal map of Nemoria glaucomarginaria. 


Prepupa essentially a stout, compacted fifth instar larva. Dorsolateral projections grad- 
ually absorbed during 3- to 4-day prepupal period. 

Pupa (Figs. 6, 7) unadorned except for light sculpturing about face and antennal bases. 
Wing cases extend to middle of A5; antennae and T3 legs often extend just beyond. Figs. 
6 and 7 show range of observed variation in relative lengths of wing and antennal cases. 
Faint venation visible on wing cases. Single setae at bases of antennae, between antennae 
and eyes, on T1 anterior to spiracle and dorsally; 2 setae on T2; seta on each abdominal 
segment in a single dorsal row and spiracular group of 3; a pair on A6 ventrally. Cre- 
master usually with 8 hooks, longest distally; 7 or 6 in some individuals. Yellow at molt, 
coloring to tan; black markings dorsal to spiracles, surrounding prespiracular setae, and 
dotted on wing veins. Mid-dorsal line dark. Eyes darken after 6-8 days, wings color to 
green with lines visible in 10-15 days; abdominal spots visible 12-24 h before eclosion. 
Emergence at dawn or dusk; pupal duration 12-18 days, with males emerging first 
(protandry). Winter diapause probably occurs as pupa, but this has not been verified, 
and at least some Nemoriini and Synchlorini diapause in third instar. 

Pattern markings on larvae remarkably constant, although degree of pattern expression 
varied slightly, and coloration varied considerably. Larvae reared during summer on 
Quercus suber L. (Fagaceae), a Mediterranean species with grayish foliage, showed little 
pattern, resulting in mostly gray-green larvae. Those reared in spring on Q. lobata Nee., 


310 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a native species, showed greatest contrast, resulting in green and brown mottled larvae. 
Whether this effect is genetic, seasonal, or a reaction to foodplant is unknown. 

Host plants. No wild collected larvae are known. All larvae were reared on new shoots 
of Quercus spp. available in Davis, California. Quercus agrifolia Nee., Q. lobata, and 
Q. suber were readily accepted. I often collected adults in association with Q. kelloggi 
Newb. in numerous localities in the Sierra Nevada and San Bernardino Mts. Material 
from Los Molinos, Tehama Co., in the northern Central Valley, probably fed on Q. 
lobata, the only oak species native to that area (Griffin & Critchfield 1976). Those col- 
lected at Norden, Nevada Co., California probably use Q. vaccinifolia Kell., the only 
oak in that area (A. M. Shapiro, pers. comm.). Those from coastal Orange and San Diego 
counties must have fed on Q. agrifolia, or on an ornamental species, for the same reason 
(Griffin & Critchfield 1976, Munz & Keck 1968). Five larvae fed Salix hindsiana Benth. 
(Salicaceae; a species acceptable to darwiniata) died in third instar. 

Behavior. All instars exhibited the often described “shaking” behavior, imitating a 
piece of dried plant material in a light breeze. This behavior is common to all Nemoriini 
and Synchlorini reared to date, but is apparently absent in the Hemitheini. First and 
second instars were observed with bits of food and fecal material attached to the body, 
a common behavior of the Synchlorini. Closer examination revealed that this adhered 
due to high humidity in the rearing containers, and not by silken threads as in the 
Synchlorini. The larvae did not replace removed material, and larvae reared in drier 
containers showed no attached materials. Young larvae, and to some extent even mature 
larvae, showed a marked tendency to remain feeding in the same place even if the food 
began to deteriorate. This behavior appears to be adaptive; dead plant material is typi- 
cally associated with caterpillar herbivory, and larvae may enhance crypsis by remaining 
at the feeding site. 

Distribution. Records are scattered throughout California west of the Great Basin (Fig. 
11). The species is also distributed northward through the Coast Ranges and Cascades to 
British Columbia (Ferguson 1985). The dense cluster of points in S California, the San 
Francisco Bay area, and in the Sierra Nevada from El Dorado to Plumas counties are no 
doubt attributable to differences in collecting intensity. I am responsible for much of the 
cluster W of Lake Tahoe. The only dubious record shown is of two specimens in the 
Bohart Museum from Alturas, Modoc Co., California, in the NE corner of the state. 
These were part of a large light trap sample at the Agricultural Inspection Station (J. S. 
Buckett, pers. comm.). There are no native oak species within 60 km of that locality 
(Griffin & Critchfield 1976), implying that the specimens were probably either carried 
in on a vehicle or mislabelled. 

Phenology. Glaucomarginaria has 2 generations annually in northern California at 
low to mid elevations, one flying in mid-May and one in early July. These dates are 
variable depending on elevation and season. Ferguson (1985) reports slightly earlier dates 
in southern California. Adults of the spring brood may be collected when the leaves of 
the foodplant begin to unfurl. I have only encountered one generation at Norden, Nevada 
Co., elevation 2,140 m. 


Nemoria darwiniata punctularia Barnes & McDunnough 


Color pattern of the mature larva of N. darwiniata punctularia Barnes & Mc- 
Dunnough is shown in Fig. 9. This species has been figured (Comstock & Henne 1940, 
as pistacearia), but the reproduction lacks details for specific identification. Nemoria 
darwiniata punctularia is usually paler than glaucomarginaria, typically yellowish green, 
with medium brown markings and whitish lines. The body is slightly stouter, and pro- 
jections correspondingly shorter. The spicules tend to be smaller and more even in size, 
giving a more velvety appearance than glaucomarginaria. The differences, however, are 
so slight that I use a tracing of Fig. 1 (glaucomarginaria) to illustrate the d. punctularia 
pattern. Fig. 9 also agrees with Dyar’s (1904) written description of d. darwiniata (Dyar) 
from British Columbia. The third instar has the same pattern of markings, but is brown 
from the lateral line dorsally to beneath the protuberances, giving the aspect of a serrate 
brown line on the side of the body. 


VOLUME 40, NUMBER 4 S| 











CALIFORNIA INSECT SURVEY 
Department of Entomology and Parasitology 


UNIVERSITY OF CALIFORNIA 


ORAFT 1955 Dasen 
115 il4 











Fic. 11. Distribution of Nemoria glaucomarginaria records in California. 


These two species are most distinguishable by pattern. In general, 
the anterior of each abdominal segment is most contrasting in glau- 
comarginaria, while the posterior of each segment is most contrasting 
in darwiniata. Glaucomarginaria larvae appear to be indistinguishable 
in color pattern and morphology from Nemoria festaria from the 
mountains of S Arizona to Texas. Female genitalia are also very similar 
between these two species (Ferguson 1985). Darwiniata is apparently 
at least oligophagous, being reared from Salix (Salicaceae) (Dyar 1904, 
pers. obs.), Quercus (Fagaceae) (Comstock & Henne 1940), and Arc- 
tostaphylos (Ericaceae) (N. McFarland notes in Los Angeles Co. Mu- 
seum of Natural History) in the laboratory. I have also recorded it in 


312 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the field from Ceanothus cordulatus Kell. (Rhamnaceae) at Alpine 
Meadows, Placer Co., California (subsequently reared ex ova on this 
host in the laboratory). 

Most larvae of the genus Nemoria examined agree in detail with the 
setal map of Fig. 10, although size and shape of projections and pro- 
tuberances differ in minor details. Thus, the above morphological de- 
scription of N. glaucomarginaria serves as a working description for 
the genus. The only exceptions known are discussed in the key below. 


DISCUSSION 


Ferguson (1985) published the most accurate figure of a Nemoria 
larva to date. A clear figure of Dichorda was given by Comstock (1960). 
No figures of Chlorosea and Phrudocentra Warren (Nemoriini), or 
Dichordophora (Dichordophorini) have been published. 

Nemoriine larval descriptions in most early publications focus on the 
bizarre morphological adaptations for crypsis and are of little value for 
taxonomic purposes at the species level. Moreover, because of the dif- 
ficulties in identifying adults before Ferguson’s (1969) revision, de- 
scriptions for most taxa need to be verified, and host records confirmed. 
As early workers did not have a broad range of species for comparative 
purposes, early figures such as those of Dammers are often misleading, 
as for example, his figure of N. leptalea (=delicataria Dyar) compared 
to the photograph in the same publication (Comstock 1960). His figure 
of N. pulcherrima (=naidaria Swett) is stylized as well (Comstock & 
Dammers 19387). Dethier (1942) figured the ultimate instar of N. ru- 
brifrontaria (Packard) with long primary setae (of which some appear 
to be missing), a condition common to the Synchlorini, but unknown 
elsewhere in the Nemoriini. Rindge (1949) used Dethier’s description 
as a basis of comparison for a preliminary generic description of Chlo- 
rosea which is correspondingly inaccurate. 

The following key will serve to rectify some of these errors. It is of 
course preliminary in that it is not based on all the species, but it will 
serve to sort out some of the characters for a future phylogenetic anal- 
ysis of the group. 


Preliminary Key to Genera of Nemoriine Larvae 


1. Dorsolateral projections on A2-6 wide, rectangular, and lamellate; setae on the 
outer corners (L1 & L2) and another (SD2) on the anterior edge of each plate 


“eesncaeBel patel WITT, NNDHE A= tac Tae Oe Ha Oe Oe ee SCL ee Dichorda 
1’. Dorsolateral projections on A2-6 conical, not wide and platelike as above _...... 2 
2. Dorsolateral projections on A2-4 tridentate, bearing three setae (SD2, L1, L2); 

the middle seta (L2) extending farthest from the body ...... Phrudocentra* 


* Based on the description of Phrudocentra in Ferguson (1985), as I have not seen larvae of this genus. Dichordophora 
(Dichordophorini) also keys out here. These two genera may be separated by locality if adults are not available: 


VOLUME 40, NUMBER 4 oe 


2'. Dorsolateral projections bidentate, bearing two setae ccc cwccccvccecvnntninientnee 8 

3. Dorsolateral projections on A2—4 with posterior seta (L1) extending farthest from 
body; mid-dorsal projections well developed, bearing both D1 setae . Chlorosea** 

3’. Dorsolateral projections on A2—4 with anterior seta (L2) extending farthest from 
body; mid-dorsal projections usually poorly developed of lacking ......... Nemoria*** 


ACKNOWLEDGMENTS 


Noel McFarland made this study possible with the loan of immature Geometrinae. R. 
Robertson and J. P. Donahue provided gravid females for rearings. A. M. Shapiro pro- 
vided help with rearings and constructive support. R. O. Schuster of the Bohart Museum, 
J. P. Donahue of the Los Angeles County Museum, W. T. Davies of the California 
Academy of Sciences, and T. Eichlin of the California Department of Food and Agri- 
culture provided access to museum specimens. D. C. Ferguson, E. M. Jakob, N. Mc- 
Farland, W. C. McGuffin, A. M. Shapiro, and the late J. S. Buckett commented on an 
earlier draft of the manuscript. This research was partially supported by a Jastro-Shields 
Research Scholarship from UC Davis. 


LITERATURE CITED 


CoMSTOCK, J. A. 1960. Inherent and applied camouflage in the subfamily Geometrinae 
(Lepidoptera) including three new life history studies. Trans. San Diego Soc. Nat. 
Hist. 12:421-439. 

ComMsTOCK, J. A. & C. M. DAMMERS. 1937. Notes on the early stages of three California 
moths. Bull. So. California Acad. Sci. 36:68-78. 

CoMSTOCK, J. A. & C. HENNE. 1940. Notes on the early stages of Nemoria pistacearia 
Pack. Bull. So. California Acad. Sci. 39:78-80. 

DETHIER, V. G. 1942. Notes on the life histories of five common Geometridae. Can. 
Entomol. 74:225-234. 

Dyar, H. G. 1904. Life histories of North American Geometridae. LVII. Psyche 11: 
12). 

FERGUSON, D. C. 1969. A revision of the moths of the subfamily Geometrinae of 
America north of Mexico (Insecta: Lepidoptera). Bull. Peabody Mus. Nat. Hist. (Yale 
Univ.) 29:1-251. 

1985. In Dominick, R. B., et al., The moths of America north of Mexico. Fasc. 
18.1. Geometroidea: Geometridae (in part). Wedge Entomological Research Foun- 
dation, Washington, D.C. 151 pp. 

GRIFFIN, J. R. & W. B. CRITCHFIELD. 1976. The distribution of forest trees in Califor- 
nia. USDA Forest Service Research Paper PSW 82, 1972 (reprinted with supple- 
ment). Pacific Southwest Forest & Range Experiment Station, Berkeley, California. 
118 pp. 

McGuFFIN, W. C. 1964. Setal patterns of the anterior abdominal segments of larvae 
of the Geometridae (Lepidoptera). Can. Entomol. 96: 841-849. 

1967. Guide to the Geometridae of Canada (Lepidoptera). I. Subfamily Ster- 

rhinae. Mem. Entomol. Soc. Can. 50:1-67. 

1977. Guide to the Geometridae of Canada (Lepidoptera). II. Subfamily En- 
nominae. 2. Mem. Entomol. Soc. Can. 101:1-191. 

Munz, P. A. & D. D. KEcK. 1968. A California flora with supplement. Univ. of Cali- 
fornia Press, Berkeley. 1681 + 224 pp. 











Dichordophora is a denizen of the desert areas of Mexico and SW United States, while Phrudocentra is from American 
subtropical and tropical regions. 

** Nemoria pulcherrima also keys out here. The adult shows affinities to both Chlorosea and Nemoria, as well as 
some unique morphological and phenological characters (Ferguson 1985). I leave it here until more data pertaining to 
its taxonomic status can be gathered. 

*** The single undetermined Nemoria larva mentioned earlier has a well developed mid-dorsal protuberance, in 
contrast to all other Nemoria examined (except pulcherrima). This may prove a phylogenetically useful character when 
more larvae have been described. 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


RINDGE, F. H. 1949. Observations on the life history of Chlorosea banksaria Sperry 
(Lepidoptera: Geometridae). Pan-Pac. Entomol. 25:24-26. 

SALKELD, E. H. 1983. A catalogue of the eggs of some Canadian Geometridae (Lepi- 
doptera), with comments. Mem. Entomol. Soc. Can. 126:1-271. 


Received for publication 31 January 1986; accepted 18 July 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 314 


GENERAL NOTE 


MASS EMERGENCES OF THE PINE WHITE, NEOPHASIA MENAPIA MENAPIA 
(FELDER & FELDER), INCOLORADO (PIERIDAE) 


Ferris and Brown (1981, Butterflies of the Rocky Mountain states, University of Okla- 
homa Press, Norman, 442 pp.) cite Neophasia menapia menapia (Felder & Felder) as 
an occasional economic pest in western montane forests, primarily of ponderosa pine 
(Pinus ponderosa Laws.) and lodgepole pine (P. contorta var. latifolia Engelmann). I 
observed mass emergences of this species in 1983 and 1984 in the pinon-juniper forest 
of western Colorado. The location was Eagle Co., White River National Forest, Frying 
Pan River Valley, 8.85 km E Basalt, ca. 2,196 m elev. In both years the emergence lasted 
3 days; 18-15 August 1983 and 3-5 August 1984. The emergences were sudden, syn- 
chronized, truly massive in nature, and all the adults disappeared as abruptly as they 
appeared. 

Earliest flight on calm, sunny mornings was recorded ca. 0745 h, and it was that of 
males typically searching for females around the high outer branch tips of mature pinon 
pine. Not until ca. 0900-0930 h did flight get fully underway, giving one the sense of 
the forest being alive with butterflies. 

Feeding was never observed in spite of special efforts to confirm Ferris and Brown's 
observation of early morning nectaring at flowers, which were abundantly available 
throughout the area. The pinon-juniper association showed no evidence of damage by 
larvae, densities of which can only be imagined. Both tree species were growing vigor- 
ously, and the pinon were filled with developing green cones. Considering their apparent 
three-day life span and the lack of feeding, it seems possible that these adults are non- 
feeders. 

Male-to-female ratio of specimens collected was 5:1 (n = 58 in 1983, 72 in 1984), 
though I believe that to be distorted by my efforts to locate the rare females. A ratio of 
50:1 is probably closer to the true situation. Females collected were so heavily gravid 
they struggled to fly. 

In the 10 years I lived at this locality (1976-85) this population explosion of N. m. 
menapia occurred only during the aforementioned 2 years. 


RONALD M. YOUNG, Systematics Research Collections, University of Nebraska State 
Museum, W436 Nebraska Hall, Lincoln, Nebraska 68588-0514. 


Received for publication 30 June 1986; accepted 7 October 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 315-317 


SOD WEBWORMS: THE LARVA OF 
MICROCRAMBUS ELEGANS (CLEM.) 
(PYRALIDAE: CRAMBINAE) 


SUZANNE ALLYSON 


Biosystematics Research Center, Agriculture Canada, 
Ottawa, Ontario, Canada KIA OC6 


ABSTRACT. Diagnostic characters of Microcrambus elegans (Clem.) larvae are giv- 
en. The long flight period in S North America suggests there are several generations 
annually; in the north there appears to be only one. The larvae were reared artificially 
on corn silk. 


Larvae of Crambinae are primarily grass feeders; some species se- 
verely damage lawns, meadows, and pastures. They have also been 
recorded damaging corn, oats, and wheat. Larvae of some species also 
feed on moss. Collections of adults are being made in E Ontario and 
W Quebec to obtain gravid females and to rear larvae from their eggs. 

There are nine North American species of Microcrambus. None of 
the larvae have been described. Microcrambus elegans is the com- 
monest and most widespread North American species of the genus. It 
occurs from Quebec S to Florida, and W to Kansas and Mississippi. In 
the northern part of its range, it flies from mid-June to late August, 
and probably only one generation occurs. In the south, it flies from 
March at least until October; there are probably several generations. 
M. elegans flies in late afternoon, early evening, and at night, coming 
commonly to light after dark. Adults appear in July and the first half 
of August in W Quebec and E Ontario. 

Moths were captured by flushing them from grass. Females were 
confined in 28 ml (7 dram) 2.9 x 6.5 cm plastic vials with snap caps 
for oviposition. Strips (1.0 x 5.0 cm) of lightly moistened blotting 
paper were placed in vials to supply drinking water and humidity. 
Females readily oviposited in these containers. Adults remained alive 
for up to one week, and, in most cases, oviposited several times, de- 
pending on the freshness of the female, laying between 20-40 eggs 
each time. Eggs were laid singly, not covered by a sticky substance, 
and were easily transferred to 4.5 x 4.5 x 1.8 cm clear plastic boxes 
with tight fitting lids. The bottom of each box was covered with blot- 
ting paper which was lightly moistened daily using a spray bottle, and 
the eggs were examined at this time for visible changes. About 50 eggs 
were placed in each rearing container. The boxes were kept out of the 
sun. The eggs had an incubation period of seven to nine days at room 
temperature. The eggs are oval, creamy white when first laid, gradu- 
ally turning yellow-orange at eclosion. Taxonomic and morphological 


316 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fics. 1-9. Larva of Microcrambus elegans, except where noted. 1, Lateral view of 
head, prothorax and mesothorax; 2, Lateral view of abdominal segment 4; 3, Lateral 
view of abdominal segments 8, 9 and 10; 4, Dorsal view of meso- and metathorax showing 
extra dorsal plate without setae; 5, Dorsal view of abdominal segments 2-8 showing 
fusion of pinacula; 6, Triordinal crochets; 7, Ventral view of abdominal segment 10 of 
pyraustine larva showing setae V1; 8, Ventral view of abdominal segment 10 of crambine 
larva showing setae V1; 9, Anterodorsal view of head. 


characters of the eggs as well as scanning electron micrographs of 
chorion sculpturing were given by Matheny and Heinrich (1972). When 
changes in the egg indicated that eclosion was about to occur, food 
was provided. Just enough food to cover the bottom of the rearing box 
was best; otherwise the first instars were difficult to locate. 

Females from which eggs were obtained were deposited as voucher 
specimens in the Canadian National Collection (CNC). Some of the 
larvae were reared to adulthood so that male genitalia could be ex- 
amined for positive identification. In general, however, the moth can 
be recognized by the brown horseshoe-shaped mark on the center of 
its folded wings. For detailed description and illustrations of the adult, 
see Felt (1894), Klots (1968), and Mauston (1970). 

Larvae of M. elegans were successfully reared on corn silk, which, 
besides being nutritious, remains fresh longer than cut grass, and is 
tender enough for young larvae to bite. The clear hue of corn silk has 
an advantage over grass when it becomes necessary to find the first 


VOLUME 40, NUMBER 4 les 


instars, which are less than 0.5 mm long. The species of grass with 
which M. elegans is normally associated are not known. 


DESCRIPTION 


Hinton’s (1949) system is used for the setae. Larvae of Crambinae can be recognized 
by the following characters: two prespiracular setae on prothorax (Fig. 1); a single trans- 
verse plate without setae posterior to dorsal pinacula on mesothorax and metathorax (Fig. 
1); only one lateral seta on abdominal (A) segment 9 (Fig. 3); setae V1 on A10 half as 
far apart as on AQ (Fig. 8), as far apart or more than in most Pyralidae (Fig. 7); crochets 
triordinal, forming a complete circle (Fig. 6). 

The larvae of M. elegans can be distinguished from other Crambinae by having 
pinacula D1 fused on dorsum of Al to 8, and pinacula D2 fused on dorsum of A2 to 8 
(Fig. 5), usually fused on dorsum of Al, but not always. The following description of last 
instar M. elegans includes only essential characters. Length 8-10 mm (N = 15). Head 
yellowish brown without markings. Stemmatal area black. Head setae as illustrated (Fig. 
9). Body pale. Pinacula distinct, light brown. Prothoracic shield brown, with scattered 
dark spots. An extra dorsal plate without setae on mesothorax and metathorax (Figs. 1, 
4). Two extra lateral plates without setae on mesothorax and metathorax (Fig. 1). Pinacula 
D1 fused on dorsum of Al to 8 and pinacula D2 fused on dorsum of A2 to 8 (Fig. 5), 
usually fused on dorsum of Al, but not always. Pinaculum SD1 surrounding spiracle on 
A2 to 7 (Fig. 2). Two extra lateral plates without setae on Al to 7 (Fig. 2), only one on 
A8 (Fig. 3). A black pit posterior to spiracle on proleg-bearing segments, probably in- 
dicative of tonofibrillary platelet (Fig. 2). 

Material examined. Quebec: Lac-Ste-Marie, female collected 25 July 1984, 3 mature 
larvae, rearing SA-84-27, CNC Voucher 68. Ontario: 10 km W Richmond, female col- 
lected 1 Aug. 1984, 12 mature larvae, rearing SA-84-34, CNC Voucher 69. 


LITERATURE CITED 


FELT, E. P. 1894. On certain grass-eating insects. Cornell Univ. Agr. Exp. Sta. Bull. 
64:47-102. 

HINTON, H. E. 1949. On the homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Entomol. Soc. 
London 97:1-37. 

Kiots, A. B. 1968. The North American Microcrambus (Lepidoptera: Pyralididae). J. 
New York Entomol. Soc. 76:9-21. 

MATHENY, E. L. & E. A. HEINRICH. 1972. Chorion characteristics of sod webworm 
eggs. Ann. Entomol. Soc. Am. 65:238-246. 

Mauston, G. W. 1970. The Crambinae of North Dakota with descriptions of some 
previously undescribed immature forms. Univ. of North Dakota, Grand Forks. 173 pp. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 318-321 


ULTRASTRUCTURE OF THE EGG OF THE 
AZALEA CATERPILLAR, DATANA MAJOR 
GROTE & ROBINSON (NOTODONTIDAE) 


GARY L. MILLER AND MICHAEL L. WILLIAMS 


Department of Entomology, Alabama Agricultural Experiment Station, 
Auburn University, Alabama 36849 


ABSTRACT. Ultrastructure of the eggs of Datana major was studied by scanning 
electron microscopy. Eggs are spheroid (1.03 x 0.78 mm) but slightly broadened toward 
the base. The chorion surface is highly ornate with a rosette-mosaic pattern surrounding 
the micropyles. The lateral surface is covered with polygonal areas having aeropyles at 
the junctions of the ridges surrounding these areas. The presence of three micropyles 
arranged deltoidally appears to be unique among the Notodontidae. 


Due to their importance as both a nursery crop and a landscape 
plant (Hill et al. 1985), greater attention has been placed on the insect 
associates of indica azaleas (Rhododendron indica (L.)). One serious 
defoliating pest of indica azaleas is the azalea caterpillar, Datana ma- 
jor Grote & Robinson (Notodontidae) (Williams et al. 1984). This cat- 
erpillar can be locally abundant and alarming to growers because of 
its extensive defoliation to azalea plantings. First stage larvae skeletonize 
the leaf surface while later stages defoliate the plant. 

Grote and Robinson first described adults of D. major in 1866. Ad- 
ditional literature concerning adult descriptions was summarized by 
Packard (1895), and descriptions of the eggs and larvae were sum- 
marized by Tietz (1972). Previous descriptions of the egg were brief 
(Dyar 1890, Packard 1895), and the only photograph illustrated eclosed 
eggs (Kuitert 1958). We more fully describe uneclosed eggs here. 


METHODS 


Fourteen eggs were field collected and 20 were obtained from 2 
females (10 eggs each) in a laboratory colony at Auburn University 
maintained on fresh indica azalea cuttings in Percival growth chambers 
at 30°C light and 24°C dark, with a 14L:10D photoperiod. Eggs were 
washed in a 1% sodium hypochlorite solution, rinsed in distilled water, 
air dried, mounted on stubs with double-coated cellophane tape and 
coated with gold-palladium in a Fullam vacuum sputter coater. Ex- 
ternal morphology was examined with an ISI-SS40 scanning electron 
microscope (SEM) using an accelerating voltage of 5 kV. Photographs 
were taken with Polaroid 55 film. Egg dimensions were recorded by 
using an ocular micrometer in a stereomicroscope, and egg color was 
estimated from standard color charts in the Munsell Book of Color 
(Munsell Color Co. 1976). Color estimates were made under “cool 


VOLUME 40, NUMBER 4 319 


white” fluorescent overhead lighting. Ultrastructure measurements were 
made using the SEM. Thirty-four eggs were observed under the SEM. 


RESULTS AND DISCUSSION 


Eggs of D. major are spheroid (Fig. 1), but slightly broadened to- 
ward the base. At the broadest point, egg width averaged 1.03 mm 
(1.00-1.06), and height 0.78 mm (0.76-0.80) (n = 10). Diameter across 
the top averaged 1.01 mm (1.00-1.06) (n = 10). Although similar in 
shape and appearance to the eggs of D. ranaeceps Guerin-Meneville 
(Peterson 1963), D. major eggs are wider (1.01 mm vs. 0.85 mm) and 
shorter (0.78 mm vs. 0.85 mm). 

The eggs have been described as sublustrous white (Dyar 1890) or 
uniform white with a large central black spot at the vertex (Packard 
1895). When the egg is viewed with the stereomicroscope, the chorion 
may even appear pearlaceous, a phenomenon which has been observed 
with other insect eggs (Arbogast & Byrd 1982). When viewed under 
fluorescent lighting, the eggs appear white to the naked eye, even 
before eclosion. By comparing eggs with the Munsell Book of Color, a 
more exact, standard description of the egg is achieved. Eggs were thus 
estimated to be 2.5 RP 8/2 or No. 2.5 red purple hue, color value No. 
8, and chroma No. 2. 

The black spot on the vertex mentioned by Packard (1895) is not 
readily noticeable on newly deposited eggs, but becomes more prom- 
inent with embyro development. The caterpillar chews through the 
chorion in the area of this spot to emerge from the egg. The thickness 
of the egg shell here is approximately 20 um. The egg is quite rigid, 
and resists crushing when pressed between thumb and forefinger. SEM 
images of eclosed eggs reveal that the egg shell is composed of nu- 
merous layers. The innermost layers are densely packed, while outer 
layers are not as tightly packed, and have larger air spaces. 

The chorion surface near the anterior pole forms a rosette-mosaic 
pattern around a depressed micropylar area (Fig. 2). This area contains 
three micropyles usually arranged deltoidally (Fig. 3), but sometimes 
linearly. The micropyles are approximately 2 wm in diameter. Hinton 
(1981) determined the number of micropyles for 19 species of noto- 
dontids but not for the azalea caterpillar. Our finding of three micro- 
pyles is the fewest observed to date in this family. The rosette-mosaic 
pattern on the egg surface represents follicular cell imprints (Margaritis 
1985). Each mosaic section is separated by deep grooves bordered by 
ridges (Fig. 4). These sections may be interconnected by narrow ridges 
to the surrounding ridges. As the pattern expands from the micropyle, 
the grooves begin to rise and eventually form only ridges around po- 


320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


- t 

‘ po t, 
‘ - 

Ke 
(aa 





"sh: [NS 
Fics. 1-5. 1, Eggs of Datana major; 2, Rosette-mosaic pattern around micropylar 


area; 3, Close-up of highlighted micropyles from Fig. 2; 4, Surface near anterior pole; 
5, Lateral surface of chorion with aeropyles (arrows). 





lygonal areas. These polygons are found throughout the surface of the 
egg. Aeropyles (Fig. 5) are usually found at ridge junctions, and av- 
erage 1.7 um (0.7-2.3 um) (n = 10) in diameter. The polygonous area 
between these ridges is highly reticulated. The density of aeropyles 
gradually diminishes toward the micropylar area. No aeropyles were 
observed in the rosette-mosaic area. 

Photomicrographs of the eggs of D. ranaeceps (Peterson 1963) re- 
semble those of azalea caterpillar. As with D. ranaeceps, D. major eggs 
were firmly attached to each other but could be easily removed from 
the substrate. We found shell thickness near that of D. ranaeceps (20 
um vs. 23 wm). Additionally, SEM images of the aeropyles and sur- 
rounding areas of the D. ranaeceps egg (Hinton 1981) closely resemble 


VOLUME 40, NUMBER 4 Syl 


those of our eggs. However, the number of micropyles differs between 
D. ranaeceps and D. major (4 vs. 8). 

Datana ranaeceps and D. major are obviously similar. Margaritis 
(1985) concluded that most eggshell features are species-specific, and 
might prove useful in clarifying phylogenetic and taxonomic problems. 
As our findings show, comparison of egg chorion morphology could 
indeed be used to reinforce current classification. 


ACKNOWLEDGMENTS 


We thank R. D. Cave, Dept. of Entomology, Auburn University, for his SEM assistance. 
We are also grateful to T. P. Mack, C. Sunderman, and J. T. Bradley, Auburn University, 
for their manuscript reviews. Ala. Agric. Exp. Stn. Journal No. 15-8697]. 


LITERATURE CITED 


ARBOGAST, R. T. & R. V. BYRD. 1982. The egg of the cadelle, Tenebriodes mauritan- 
icus (L.) (Coleoptera: Trogositidae): Fine structure of the chorion. Entomol. News 
93:61-66. 

Dyar, H. G. 1890. Notes on two species of Datana with descriptions of their larval 
stages. Psyche 5:114—-120. 

HILL, M. L., L. E. WILson & R. SHUMACK. 1985. Azaleas important crop in Baldwin 
and Mobile counties. Ala. Agric. Exp. Stn. Highlights of Agric. Res. 32(8):18. 

HINTON, H. E. 1981. Biology of insect eggs. Vols. 1-3. Pergamon Press Inc., New York. 
1125 pp. 

KUITERT, L. C. 1958. Insect pests of ornamental plants. Univ. Fla. Agric. Exp. Sta. Bull. 
595. 51 pp. 

MarGanritis, L. H. 1985. Structure and physiology of the eggshell, pp. 153-230. In 
Kerkut, G. A. and L. I. Gilbert (eds.), Comprehensive insect physiology biochemistry 
and pharmacology. Vol. 1. Pergamon Press Inc., New York. 

MUNSELL COLOR Co. 1976. Munsell book of color. 2 Vols. Macbeth Division of Koll- 
morgen Corp., Baltimore, Maryland. 

PACKARD, A. S. 1895. Systematic revision of the Notodontidae with special reference 
to their transformation. Mem. Natl. Acad. Sci. 7:87—284. 

PETERSON, A. 1963. Some eggs of moths among the Amatidae, Arctiidae, and Noto- 
dontidae—Lepidoptera. Fla. Entomol. 46:169-182. 

TrETZ, H. M. 1972. An index to the described life histories, early stages and hosts of 
the macrolepidoptera of the continental United States and Canada I. Allyn Museum 
of Entomology. 536 pp. 

WILLIAMS, M. L., G. L. MILLER & B. J. SHEFFER. 1984. Azalea caterpillars damage 
azalea foliage. Ala. Agric. Exp. Stn. Highlights of Agric. Res. 31(4):4. 


Received for publication 27 February 1986; accepted 24 July 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 322-326 


NEW SPECIES OF OLETHREUTINE MOTHS (TORTRICIDAE) 
FROM TEXAS AND LOUISIANA 


EDWARD C. KNUDSON 


808 Woodstock, Bellaire, Texas 77401 
Research Associate, Florida State Collection of Arthropods, 
Gainesville, Florida 32602 


ABSTRACT. Three new species are described; the male imago and male and female 
genitalia of each are figured. Eucosma rosaocellana is described from eight specimens 
from northwest Texas, and contrasted with E. salaciana Blanchard & Knudson. Dichro- 
rampha broui is described from 37 specimens from southeast Louisiana and northeast 
Texas, and is contrasted with D. leopardana (Busck) and D. incanana (Clemens). Pam- 
mene medioalbana is described from nine specimens from central Texas, and is con- 
trasted with Cydia latiuscula (Heinrich) and C. gallaesalaciana (Riley). 


The following three new species of olethreutine moths are described 
to facilitate completion of a catalogue-checklist of the moths of Texas, 
a project originated by André Blanchard. Many other undescribed 
species of Texas Lepidoptera remain or are likely to be discovered in 
the Tortricidae and other families. In many such cases, the families or 
their subdivisions are in such great need of revision that isolated de- 
scriptions of new species could confuse the situation. Since the last 
comprehensive revision of Eucosma (Heinrich 1923), numerous new 
Eucosma species have been described, but the one described below is 
quite distinctive, and is closely related to E. salaciana Blanchard & 
Knudson (1981). Heppner (1981) described two new species of Di- 
chrorampha from Florida, and Miller (1983) reduced five Dichro- 
rampha species names to synonyms, and summarized other recent work 
in this genus. No new species of Pammene have been described from 
North America since Heinrich’s (1926) revision. The holotypes of all 
species described below are in the U.S. National Museum of Natural 
History (USNM). 


Eucosma rosaocellana E. C. Knudson, new species 


(Figs. 1, 4, 7) 


Description. Head: Front brownish white; vertex pale orange brown; labial palpi 
porrect, 2nd segment equal to 1% eye diameter, brownish white dorsally, scale tuft 
orangish, shading to gray at apex, 3rd segment equal to about % eye diameter, brownish 
white; antennae brownish white, ventral setae about 4 width of segments (n = 5). Thorax: 
Orangish brown above, with 3 transverse whitish bands; whitish beneath; legs whitish, 
tibiae and tarsi with several light gray bands. Abdomen: Brownish white above and 
beneath. Forewing: Ratio of length of costal fold (in male) to forewing length 0.32 (n = 
5). Ground color whitish, reticulated with irregular vertical bands of grayish brown, and 
variably suffused with orange brown below costal margin and over lower fold. Subbasal 
fascia orange brown, margined with blackish brown and weakly represented at costal 
margin; median fascia grayish brown, margined with blackish brown and weakly rep- 
resented beyond pretornal spot. Ocelloid patch a trefoil of pinkish opalescent scales 


VOLUME 40, NUMBER 4 oe 


surmounted by a grayish brown patch containing two or three short horizontal black 
dashes. Costal margin strigulate with about 15 evenly spaced strong blackish brown 
dashes, with a like number of thinner paler dashes between them, all more or less 
connected to the irregular vertical bands. Fringe grayish brown mixed with white, mainly 
white over tornal third. Hindwing: Pale grayish brown, paler toward base; darker line 
at termen and a second dark line at the base of the whitish fringe. Length of forewing: 
Males, 7.1-10.0 mm, average 8.7 mm (n = 7); female, 8.6 mm (n = 1). Male genitalia 
(Fig. 4) (n = 8): Sacculus with ventral margin forming a rounded angle of 105-110 
degrees at neck constriction; cucullus broadly rounded at apex, produced ventrally, with 
many short, spinose setae; cucullus width to length ratio 1.9-2.1; vesica with about 12 
deciduous cornuti. Female genitalia (Fig. 7) (n = 1): Papillae anales large, bilobed; 
apophyses anteriores twice as long as apophyses posteriores; lamina postvaginalis well 
sclerotized, tapering toward an arcuate posterior margin; lamina antevaginalis broad, 
weakly sclerotized; ductus bursae slightly dilated over middle 4%, sclerotized on dorsolat- 
eral surface, ductus seminalis from just below middle; corpus bursae ovoid, tapering 
anteriorly, membranous, with two equal sized thornlike signa. 

Types. Holotype (Fig. 1): Male, Texas, Hemphill Co., Gene Howe Wildlife Manage- 
ment Area, 18-V-85, collected by E. C. Knudson. Paratypes: Same data and collector as 
holotype, 6 males (three with genitalia on slides ECK 1159, 1374, 1375), retained by 
author; Cottle Co., Matador Wildlife Management Area, 30-VII-83, 1 female (genitalia 
on slide ECK 1167), in USNM. 


Remarks. This species is closely related to Eucosma salaciana Blan- 
chard & Knudson (1981), which has a similar forewing pattern, but is 
marked with dark gray, lacking any trace of orange, and is smaller, 
with an average forewing length in males of 7.1 mm (n = 28). Male 
genitalia of the two species are very similar, but the length/width ratio 
of the cucullus in salaciana is 2.2-2.4, and the ventral angle of the 
sacculus is 90 degrees (n = 4). Female genitalia of E. salaciana differ 
in the lamina postvaginalis, which is bilobed posteriorly, and in the 
apophyses posteriores, which are one-third the length of the apophyses 
anteriores (n = 2). The name of the new species refers to the pinkish 
ocelloid patch. 


Dichrorampha broui E. C. Knudson, new species 


(Figs. 2, 5, 8) 


Description. Head: Front and vertex gray; labial palpi exceeding front by 2 eye 
diameters, dark gray with an oblique whitish band on lateral surface of 2nd segment; 
antennae gray. Thorax: Gray above, paler beneath; legs whitish, tibiae and tarsi banded 
with gray and white. Abdomen: Shiny yellowish gray above and beneath. Forewing: 
Male without costal fold. Ground color grayish white, marked with numerous closely 
spaced blackish strigulations, outwardly oblique from dorsal margin to radial vein, from 
there angled inwardly to costal margin. Basal and median fasciae weakly indicated by 
stronger, darker strigulations. Before termen at middle are three short black horizontal 
dashes, at tornus, a black dot separated from the three dashes by a whitish patch. Termen 
slightly produced at apex. On distal half of costal margin are five equally spaced white 
spots, the inner two or three geminate. Terminal line dark gray; fringe shining yellowish 
gray. Hindwing: Light gray, darker near termen; terminal line dark gray; fringe shining 
yellowish gray. Length of forewing: Males, 5.2-6.0 mm, average 5.5 mm (n = 10); 
females, 4.9-6.1 mm, average 5.6 mm (n = 10). Male genitalia (Fig. 5) (n = 4): Valva 
with costa bent anteriorly; neck incurvation broad, tapering to a narrow constriction 
before cucullus; cucullus ovoid, entirely anterior and extending well basad of neck in- 


324 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





j 7 “= 8 9 

Fics. 1, 4,7. Eucosma rosaocellana. 1, Holotype male; 4, Male genitalia; 7, Female 
genitalia. 

Fics. 2, 5, 8. Dichrorampha broui. 2, Holotype male; 5, Male genitalia; 8, Female 
genitalia. 

Fics. 3, 6,9. Pammene medioalbana. 3, Holotype male; 6, Male genitalia; 9, Female 
genitalia. 

Scale lines represent 1 mm. 


VOLUME 40, NUMBER 4 SAD) 


sertion; vesica with numerous tiny deciduous cornuti. Female genitalia (Fig. 8) (n = 2): 
Apophyses posteriores % the length of apophyses anteriores; ostium bursae sclerotized, 
bilobed, with a short median groove; ductus bursae sclerotized over posterior half, ductus 
seminalis from near middle of anterior half; corpus bursae membranous with 1 thornlike 
signum. 

Types. Holotype (Fig. 2): Male, Louisiana, St. Tammany Parish, 4.2 miles (6.7 km) 
NE Abita Springs, 12-IV-85, collected by V. A. Brou. Paratypes: Same locality and 
collector as holotype, 20-IV-84, 1 male, 1 female; 21-IV-84, 2 females; 22-IV-84, 1 male, 
1 female; 29-IV-84, 1 female; 2-V-84, 1 female; 3-V-84, 1 male (genitalia on slide USNM 
25933 by J. F. Gates Clarke in USNM); 7-IV-85, 1 female; 12-IV-85, 4 males, 2 females; 
13-IV-85, 1 male, 1 female; 18-IV-85, 1 male (genitalia on slide ECK 1302); 19-IV-85, 2 
males (one with genitalia on slide ECK 1378), 1 female; 20-IV-85, 4 males, 1 female; 23- 
IV-85, 1 male; 24-IV-85, 1 female; 27-IV-85, 1 female; all collected by V. A. Brou. Texas, 
Morris Co., Daingerfield State Park, 22-IV-85, 1 male (genitalia on slide ECK 1136), 1 
female (genitalia on slide WEM 25852 by W. E. Miller); Jasper Co., Martin Dies Jr. State 
Park, 27-IV-86, 2 males, 2 females (one with genitalia on slide ECK 1384), collected by 
E. C. Knudson. Four Louisiana paratypes in USNM; four retained by author; remainder 
retained by collector. One Texas paratype in USNM, remainder retained by author. 


Remarks. This species is similar in both male and female genitalia 
to Dichrorampha leopardana (Busck) (as illustrated by Heinrich 1926), 
but broui can easily be distinguished by forewing pattern, which lacks 
the orange striping of leopardana. Male genitalia of D. broui are also 
similar to those of D. incanana (Clemens) (as represented by Heinrich 
1926), but in that species the male has a costal fold. The new species 
is named for its discoverer, Vernon A. Brou. 


Pammene medioalbana E. C. Knudson, new species 


(Figs. 3, 6, 9) 


Description. Head: Front and vertex light gray; labial palpi light gray, exceeding front 
by % eye diameter; antennae dark gray. Thorax: Above, anterior % dark gray, posterior 
¥% whitish; whitish beneath; legs whitish, tarsi faintly banded with light gray. Abdomen: 
Whitish gray above and beneath; male with specialized sex scales on 6th and 7th terga, 
consisting of 5 or 6 irregular rows of persistent scale tufts on 6th tergum, and 2 or 3 rows 
of weak scale tufts on 7th tergum; both obscured by superficial scaling. Forewing: Ground 
color white; basal 4% suffused with dark brown scales, forming a dark basal patch; median 
area mainly white with 3 obscure, darker, oblique striations; subterminal area from dorsal 
margin % the distance from base to just before apex dark brown with a faint coppery 
luster, enclosing 4 or 5 short, horizontal black dashes margined by narrow silvery metallic 
bars; costal margin whitish with short, dark brown strigulations, most distinct over apical 
¥3; terminal! line black, fringe shining gray. Hindwing: Dark gray, fringe light gray. 
Length of forewing: Males, 4.0—-4.2 mm, average 4.1 mm (n = 6); females, 4.2-4.4 mm, 
average 4.3 mm (n = 3). Male genitalia (Fig. 6) (n = 3): Sacculus with a few strong 
hairlike setae before slight neck incurvation; cucullus broadly rounded; vesica with 3 or 
4 small spinelike nondeciduous cornuti at apex, arranged in a line. Female genitalia 
(Fig. 9) (n = 2): Apophyses posteriores very slender, equal in length to broader apophyses 
anteriores; ostium bursae a short membranous funnel, lamina postvaginalis well sclero- 
tized, U-shaped; ductus bursae short, well sclerotized; corpus bursae membranous with 
two small thornlike signa. 

Types. Holotype (Fig. 3): Male, Texas, Gonzalez Co., Palmetto State Park, 23-III-85, 
collected by E. C. Knudson. Paratypes: Same data and collector as holotype, 5 males (3 
with genitalia on slides ECK 1122, 1267, and 1278), 3 females (2 with genitalia on slides 
ECK 1163 and 1385). One male and one female paratype in USNM, others retained by 
author. 


326 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Remarks. This species is more similar in forewing pattern to Cydia 
lautiuscula (Heinrich 1926) (illustrated by Miller 1976) and C. gallae- 
salaciana (Riley) (as represented by Heinrich 1926) than to other North 
American species of Pammene, but the wing venation, abdominal scale 
tufts, and male and female genitalia are characteristic of the latter 


genus. 


ACKNOWLEDGMENTS 


I thank R. L. Brown, J. F. Gates Clarke, and W. E. Miller for examining specimens 
and providing helpful suggestions. I am also grateful to Vernon A. Brou for providing 
specimens, and to the Texas Parks and Wildlife Dept. for providing access to collecting 
localities. 


LITERATURE CITED 


BLANCHARD A. & E. C. KNUDSON. 1981. Two new species of Eucosma Hiner (Tor- 
tricidae) from Texas. J. Lepid. Soc. 35:173-178. 

HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucos- 
minae of the family Olethreutidae. U.S. Natl. Mus. Bull. 123. 298 pp. 

1926. Revision of North American moths of the subfamilies Laspeyresiinae and 
Olethreutinae. U.S. Natl. Mus. Bull. 182. 216 pp. 

HEPPNER, J. B. 1981. Two new Dichrorampha (Lepidoptera: Tortricidae) from Florida. 
Fla. Entomol. 64:271-276. 

MILLER, W. E. 1976. A new species of Laspeyresia from Michigan (Lepidoptera: 
Olethreutidae). Great Lakes Entomol. 9:171-172. 

1983. New synonymies in nearctic Dichrorampha (Lepidoptera: Tortricidae). 

Proc. Entomol. Soc. Wash. 85:727-783. 








Received for publication 10 March 1986; accepted 13 August 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 327-346 


RESURRECTION OF CATASTEGA CLEMENS AND 
REVISION OF THE EPINOTIA VERTUMNANA (ZELLER) 
SPECIES-GROUP (TORTRICIDAE: OLETHREUTINAE) 


RICHARD L. BROWN 


Mississippi Entomological Museum, Drawer EM, 
Mississippi State, Mississippi 39762 


ABSTRACT. Catastega is recognized as a valid genus; C. timidella, the type species, 
and C. aceriella are resurrected from synonymy with Epinotia; and E. marmoreana is 
transferred to Catastega. In the Epinotia vertumnana species-group revision, E. atri- 
striga is synonymized with E. zandana, and Paedisca celtisana is synonymized with E. 
laracana. The previously misidentified E. laracana is described as a new species. Lec- 
totypes are designated for Grapholitha subnisana, a junior synonym of C. aceriella, and 
Paedisca vertumnana, which has been previously misidentified as E. zandana. Imagos 
and female genitalia of Catastega species are illustrated; imagos and male and female 
genitalia of six species in the E. vertumnana species group are described and illustrated. 


The treatment of North American Epinotia Htibner by Heinrich 
(1923) included two species groups with similarly developed male gen- 
italia, both having plesiomorphic forms of an uncus and valva. Epinotia 
vertumnana (Zeller) and five related species comprise the most gen- 
eralized species-group of the genus, whereas the E. aceriella species- 
group does not appear congeneric with the former. Although a system- 
atic study of world Epinotia is near completion, these two species- 
groups are treated here to clarify nomenclature and make names avail- 
able to other workers. 

For species descriptions, specimens and genitalia were examined 
with a phase-contrast microscope and stereomicroscope with ocular 
micrometer, which was used for all measurements. Measurements of 
forewings are accurate to 0.1; other measurements are accurate to 0.01. 
The forewing length (FWL) was measured from base to apex, includ- 
ing fringe. The length of sacculus was measured as a straight line from 
ventral base of valva to beginning of the cucullus, as restricted to the 
apical portion of the valva with subparallel dorsal and ventral margins 
(Fig. 8). The number of deciduous cornuti in the aedeagus was ob- 
tained by counting their sockets, which were usually distinct. For scan- 
ning electron microscopy, specimens were coated with gold-palladium 
for six min and examined with a Hitachi HH-S-2R scanning electron 
microscope. 

Partial synonymies are given for Catastega species to include pre- 
vious binomial combinations and major references with illustrations of 
life stages. Complete synonymies are given for species in the E. ver- 
tumnana group. Specimens in collections of the following individuals 
and institutions were examined: André Blanchard (AB), American Mu- 


328 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


seum of Natural History (AMNH), Academy of Natural Sciences, 
Philadelphia (ANSP), Bryant Mather (BM), British Museum of Natural 
History (BMNH), Canadian National Collection (CNC), Cornell Uni- 
versity (CU), Edward C. Knudson (ECK), Illinois Natural History Sur- 
vey (INHS), John G. Franclemont (JGF), J. Richard Heitzman (JRH), 
Kansas State University (KSU), Natural History Museum of Los An- 
geles (LACM), Museum of Comparative Zoology (MCZ), Mississippi 
Entomological Museum (MEM), Michigan State University (MSU), 
Richard L. Brown (RLB), University of California, Berkeley (UCB), 
United States National Museum of Natural History (USNM). Label 
data from these specimens have been recorded and deposited in the 
Mississippi Entomological Museum. 


RESURRECTION OF CATASTEGA 


In 1862 Brackenridge Clemens erected the genus Catastega for ac- 
eriella and timidella, based solely on larval habits. Larvae incorporate 
frass with silk to form serpentine tubes on the underside of leaves (Fig. 
1), similar to some Gelechiidae and Pyralidae. The tubes and feeding 
area are covered by a loose web of silk, which causes the leaf to crum- 
ple as larvae mature. The family placement was uncertain until H. G. 
Dyar reared an adult of C. timidella, which was determined to be a 
tortricid (Busck 1903). Heinrich (1923) treated Catastega as a synonym 
of Epinotia and described E. marmoreana, which is transferred to 
Catastega here. 

Catastega is characterized as follows: forewing usually with well 
developed pretornal triangular spot, often with reticulations between 
fascia, costal strigulae present or absent, male costal fold present or 
absent; male genitalia with uncus bifid, socii broad, setose, ventrally 
fused with bases of gnathos, anellus not closely surrounding base of 
aedeagus, often cuplike, valva with saccular spine cluster, cucullus 
poorly defined or delimited by deep ventral invagination; female with 
lamella postvaginalis reduced, lamella antevaginalis developed and 
forming conelike sterigma around ostium, ductus bursae with sclero- 
tized band posterior to inception of ductus seminalis, two signa present. 

The previous inclusion of Catastega with Epinotia was based on 
plesiomorphic characters that also are present in some Olethreutini, 
such as Omiostola Meyrick: uncus well developed and bifid, socius 
broadly joining tegumen, valva with a saccular spine cluster. The male 
anellus and female sterigma are apomorphic for Catastega. 

Although Catastega is known to occur only in North America, the 
genus appears to be closely related to Pseudacroclita Oku, which is 
endemic to Japan. Larvae of the latter genus also spin a tubular nest 


VOLUME 40, NUMBER 4 329 





Fic. 1. Larval feeding tube (arrow) of Catastega aceriella on Acer sp., Ithaca, New 
York. 


covered by a sparse web on the lower surface of leaves. Female geni- 
talia of the type species, P. hapalaspis (Meyrick), are very similar to 
Catastega species, but the male differs in having a short, broad uncus, 
and a different form of aedeagus and valva (Oku 1979). 


Catastega Clemens, revised status 


Catastega Clemens (1861 [1862]:86). Type species: Catastega timidella Clemens, by 
subsequent designation (Busck 1903:852). 


Catastega aceriella Clemens, revised status 


(Figs. 2, 5) 


Catastega aceriella Clemens (1861 [1862]:86). 

Hedya signatana Clemens (1864 [1865]:514); Miller (1973:222, fig. 41, imago). 

Steganoptycha variana Clemens (1864 [1865]:520); Miller (1973:224, fig. 49, imago). 

Grapholitha subnisana Zeller (1875:294). 

Semasia signatana; Fernald (1902 [1903]:462); Felt (1905:168, fig. 24, larval work); Mosh- 
er (1916:54). 

Gelechia aceriella; Busck (1902 [1903]:515). 

Enarmonia aceriella; Fernald (1908:39, 56). 

Eucosma sigmatana Barnes & McDunnough (1917:172, misspelling). 

Epinotia aceriella; Heinrich (1923:244, fig. 372, 6 genitalia); MacKay (1959:118, fig. 111, 
larva), (1962:638); Miller (1973:211); Coté & Allen (1973:463-470, figs. 3-9, egg, 
larval work, pupa, imago); Johnson & Lyon (1976:178, Pl. 81, figs. a—c, larval work). 


Types. aceriella description based on larval work; type locality: unknown locality in 
North America. signatana—Holotype, 6, in ANSP; type locality: Virginia. variana— 
Lectotype, 2, designated by Miller (1973); in ANSP; type locality: Maine or Easton, 
Pennsylvania. subnisana—Lectotype here designated, 4, ““Mass.-Maine? Packard” [green 
label], genitalia slide BM No. 11597; a lectotype label was previously affixed to the 
specimen by N. S. Obraztsov, but no formal designation was published; a lectotype label 
dated 1986 with designation by R. L. Brown has been added to the specimen; in BMNH; 
type locality: “Maine-Mass.?”. 


330 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 2-4. Imagos of Catastega species. 2, C. aceriella, male, Tompkins Co., New 
York; 3, C. marmoreana, female, Coconino Co., Arizona; 4, C. timidella, male, Tomp- 
kins Co., New York. 


Distribution. Eastern North America from S Ontario and Nova Scotia to North Car- 
olina and Tennessee, W to Illinois and Minnesota. 


Hosts. Acer spp. 
Material examined. Types of signatana, variana, subnisana; 91 6, 36 2 (AMNH, 


ANSP, BMNH, CNC, CU, INHS, JGF, RLB, UCB, USNM). 


Catastega marmoreana Heinrich, new combination 


(Figs. 3, 6) 


Epinotia marmoreana Heinrich (1923:222, fig. 349, 6 genitalia). 


Type. Holotype, 6, USNM Type No. 24852, genitalia slide “CH 2 Nov. 1920”; in 
USNM. Type locality: Stockton, Utah. 
Distribution. Western United States from N Arizona and New Mexico to S Wyoming. 


Host. Unknown. 
Material examined. 6 Type, 15 6, 29 2? (AMNH, INHS, JGF, LACM, USNM). 


Catastega timidella Clemens, revised status 


(Figs. 4, 7) 


Catastega timidella Clemens (1861 [1862]:96). 

Gelechia timidella; Busck (1902 [1903]:518). 

Enarmonia timidella; Fernald (1908:39). 

Epinotia timidella; Heinrich (1923:223, fig. 373, 6 genitalia); MacKay (1962:638, fig. 7, 
larva); Miller (1973:224). 


Type. timidella description based on larval work; type locality: St. Paul, Minnesota? 
Distribution. Eastern North America from S Canada to Virginia, W to Illinois and 
Minnesota; British Columbia (probable introduction). 


Hosts. Quercus spp. 
Material examined. 71 6, 50 2 (AMNH, ANSP, CNC, CU, INHS, JGF, LACM, RLB, 
UCB, USNM). 


Discussion 


The three described species are differentiated easily by wing pattern. 
Catastega timidella and C. aceriella cannot be separated easily by 
characters of male genitalia. The female C. aceriella has sternum VII 
without rugae, tergum VIII and papillae anales narrower, and sterigma 


VOLUME 40, NUMBER 4 331 





Fic. 5. Female genitalia of Catastega aceriella, Pittsburgh, Pennsylvania, USNM 
slide 17600, with sterigma and ductus bursae enlarged. Scale line = 1 mm. 


more rounded than C. timidella. Both sexes of the western C. mar- 
moreana differ greatly from eastern species, especially in shape of 
male valva and female sterigma. More than 10 undescribed species 
occurring in SW United States and Mexico and at least one new species 
in E United States have been collected recently. These will be de- 
scribed when additional material becomes available; some are repre- 
sented by only one sex. 


332 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


| 
Ure 
is 


Onn 

Sit 

SR 
SAN) 


\ 
NY) is 





Fics. 6,7. Female genitalia of Catastega species. 6, C. marmoreana, Colorado Springs, 
Colorado, RLB slide 471 (INHS); 7, C. timidella, Aweme, Manitoba, RLB slide 523 
(CNC). Scale line = 1 mm. 


REVISION OF THE EPINOTIA VERTUMNANA SPECIES-GROUP 


The E. vertumnana species-group of Eucosmini includes six species 
of grayish brown moths, all of which occur E of the Rocky Mountains 
in North America. Species in this group are the most difficult to iden- 
tify among all Epinotia because of obscure and variable forewing fas- 
ciae in most, and similarity of color pattern and male genitalia among 


VOLUME 40, NUMBER 4 333 











Fic. 8. Male genitalia of Epinotia vertumnana, Dallas, Texas, RLB slide 1155 (MCZ). 
DAP, dorsal anellar plate; length of sacculus measured as straight line between arrows. 
Scale line = 1 mm. 


some. Of the six treated here, five were misidentified in the last revision 
of North American Eucosmini (Heinrich 1928). 

Imagos of this group are among the first tortricids to fly during early 
spring, and are often collected with Pseudexentera and Chimoptesis. 
Epinotia can be separated from the latter two genera and other Eu- 
cosmini by the following genital characters: in the male, uncus devel- 
oped, socius arising laterally rather than dorsally from tegumen, gna- 
thos usually not fused medially, anellus usually closely surrounding 
aedeagus and extended dorsally as an anellar plate; and in the female, 
sternum VII not fused with sterigma and not posteriorly invaginated 
around ostium, lamella antevaginalis reduced, and ductus bursae with 
sclerotized, denticulate band or plate near inception of ductus semin- 
alis. 

Species in the vertumnana group share the following characters: 
length of third segment of labial palpus less than half the length of 
second; scales of head, tegulae, and mesonotum concolorous with fore- 
wing ground color; forewing outer margin straight, forming an acute 
angle with costal margin; male costal fold present or absent, enclosing 
hair pencils; hindwing light grayish brown, without contrasting colors; 
in the male, uncus bifid from near base; socius arising from bulbous 
expansion of tegumen, elongate, apically rounded, setose on dorsal 


334 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


margin from apex to base of uncus; gnathos arising from tegumen and 
base of socius, heavily sclerotized basally; aedeagus moderately long 
and stout; anellus closely surrounding aedeagus basally, divergent api- 
cally; caulis of juxta V-shaped in cross-section; valva with well defined 
saccular spine cluster; cucullus not well defined nor delimited by neck; 
in the female, sternum VII with medial area more lightly sclerotized 
than lateral areas, without depressions and microtrichia; tergum VIII 
setose, without scales; lamella postvaginalis with microtrichia present 
medially, setose laterally; ductus bursae spiraled, medial sclerotized 
band encircling ductus bursae much wider on one side than other; two 
signa present. Species of the vertumnana group can be separated easily 
from other Epinotia by the grayish brown forewings with acute apex 
and poorly defined fasciae, the form of bifid uncus, the valva with a 
well defined saccular spine cluster and poorly defined cucullus (Fig. 
8), and in the female, by the lightly sclerotized medial line of sternum 
VII and form of the sterigma. 


Epinotia zandana (Kearfott) 
(Figs. 9, 10, 18, 19, 26) 


Eucosma zandana Kearfott (1907:25). 
Eucosma peristicta Meyrick (1912:34, invalid replacement name). 
Epinotia atristriga Clarke (1953:228). NEW SYNONYMY. 


Adult. 6 Types, zandana, atristriga; 58 6, 48 2 examined (AMNH, ANSP, CNC, CU, 
ECK, JRH, INHS, KSU, MSU, RLB, UCB, USNM). 

Forewing (Figs. 9, 10). FWL: 6.0-8.5 mm; length of costal fold 0.42-0.46 FWL (12n); 
most specimens dark grayish brown peppered with variable number of white or white- 
tipped scales, white scales usually forming a longitudinal sinuate line on inner marginal 
half; some specimens with narrow, brown, subbasal and median fasciae, median fascia 
interrupted near CuA1 to form pretornal triangular spot; some specimens with brown, 
continuous or interrupted, longitudinal streak from base to apex. 

Male genitalia (Figs. 18, 19). Width of uncus at bifurcation 0.44—-0.48 greatest width 
of juxta; dorsal anellar plate with straight lateral margins, apically curved outward; 
aedeagus with 9-11 cornuti (5n); valva abruptly narrowed beyond sacculus, cucullus 
much longer than sacculus; saccular spine cluster rounded. Twenty-three preparations 
examined (AMNH, CNC, ECK, MSU, UCB, USNM). 

Female genitalia (Fig. 26). Tergum VIII with row of setae on posterior margin, bare 
medially, anterior margin with rounded medial notch; papillae anales large, with dense 
setae and microtrichia, facing laterally, subequal in width anteriorly and posteriorly; 
length of anterior apophyses 1.0-1.1 length of posterior apophyses (7n); posterior margin 
of sterigma straight to slightly concave; bursa seminalis without spinules; width of pos- 
terior signum 1.7—2.7 width of anterior signum (8n). Twenty-one preparations examined 
(ANSP, CNC, INHS, MSU). 

Types. Eucosma zandana—Lectotype, 6, genitalia slide by Klots, 26 Oct. 1941; des- 
ignated by Klots (1942); in AMNH. Type locality: Cincinnati, Ohio. Epinotia atristriga — 
Holotype, 4, genitalia slide CH#1, 13 Aug. 1940; in USNM. Type locality: Putnam Co., 
Illinois. The zandana lectotype is similar to the specimen in Fig. 9; the atristriga holotype 
is similar to the specimen in Fig. 10. The atristriga holotype is labeled as collected on 
24 March 1938, rather than 17 March 1938, as given in error by Clarke (1953). The 
female genitalia illustrated by Clarke for atristriga are those of E. laracana. 


VOLUME 40, NUMBER 4 300 





Fics. 9-17. Imagos of Epinotia species. 9, E. zandana, female, no data; 10, E. 
zandana, male, Ottawa, Ontario; 11, E. bicordana, male, Aweme, Manitoba; 12, E. 
laracana, male, Dallas, Texas; 13, E. laracana, female, Ottawa, Ontario; 14, E. xandana, 
female, Cincinnati, Ohio; 15, E. sotipena, female paratype, Ithaca, New York; 16, E. 
vertumnana, female, Merivale, Ontario; 17, E. vertumnana, female, Clifton Springs, 
New York. 


Geographical distribution and flight times. S Ontario and Quebec (24 Mar.—29 Apr.) 
to Massachusetts (16 Mar.-1 May), W to Michigan (21 Mar.—13 Apr.), and S to Arkansas 
(4-13 Mar.) and E Texas (17 Feb.-10 Mar.). 

Host. Crataegus sp. (one specimen reared by T. N. Freeman in Ontario). 


Discussion. Specimens that Heinrich (1928) treated as E. zandana 
are conspecific with the lectotype of E. vertumnana. Most specimens 
of E. zandana can be differentiated from other members of the species 
group by the darker color of the forewings. This species is easily iden- 
tified by the broad base of uncus, rounded saccular spine cluster of 
valva, and cucullus that is much longer than the sacculus. The female 
is distinctive in having broad papillae anales with dense setae and 
microtrichia, and can be identified without dissection. The short ovi- 
positor (with posterior and anterior apophyses subequal in length) and 


336 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





VOLUME 40, NUMBER 4 O37 


the densely setose papillae anales suggest that eggs are not inserted 
into crevices or buds, in contrast to the inserting form of ovipositor 
with reduced papillae anales (Figs. 30, 31). The papillae anales of many 
females have debris packed between setae. 


Epinotia bicordana Heinrich 
(Figs. 11, 20, 27) 


Epinotia bicordana Heinrich (1928:220, fig. 368). 


Adult. 6 Type, 12 6, 7 2 examined (AMNH, CNC, USNM). 

Forewing (Fig. 11). FWL: 6-7 mm; costal fold absent; basal third of costa rolled 
dorsally and slightly posteriorly; uniformly grayish brown, some specimens with narrow, 
dark grayish brown subbasal and median fasciae. 

Male genitalia (Fig. 20). Width of uncus at bifurcation 0.24 greatest width of juxta 
(1n); dorsal anellar plate with sinuate lateral margins; aedeagus with six cornuti (ln); 
valva abruptly narrowed beyond sacculus, cucullus much longer than sacculus; saccular 
spine cluster elongate. Two preparations examined (AMNH, USNM). 

Female genitalia (Fig. 27). Tergum VIII with 1-2 rows of setae on posterior margin, 
anterior margin W-shaped, median notch acute; papillae anales large, facing ventrally, 
posterior cleft shallow, subequal in width anteriorly and posteriorly; posterior and an- 
terior apophyses subequal in length; posterior margin of lamella postvaginalis slightly 
sinuate; bursa seminalis without spinules; width of posterior signum 1.6 width of anterior 
signum. One preparation examined (CNC). 

Type. Holotype, 4, genitalia slide “#5, Jan. 30, 1920”; in AMNH. Type locality: 
Aweme, Manitoba. 

Geographical distribution and flight times. Known only from Aweme, Manitoba (26 
Mar.—27 Apr.). 

Host. Unknown. 


Discussion. This species is superficially similar to uniformly colored 
individuals of E. zandana and E. vertumnana. Males of E. bicordana 
differ from males of all related species in lacking a forewing costal 
fold. The female can be easily identified by the combination of the 
W-shaped anterior margin of tergum VIII, and the broad, ventrally 
facing papillae anales. Debris was absent on the papillae anales of the 
single female examined. 


Epinotia xandana (Kearfott) 
(Figs. 14, 21, 28) 


Paedisca vertumnana; Zeller (1875:310, part, var. c). 
Eucosma xandana Kearfott (1907:24). 
Eucosma atacta Meyrick (1912:34, invalid replacement name). 


—_— 


Fics. 18-25. Male genitalia of Epinotia species. 18, E. zandana, Putnam Co., Illinois, 
JFGC 10564 (USNM); 19, E. zandana, Cincinnati, Ohio, Klots 26 Oct. 1941, lectotype 
(AMNH); 20, E. bicordana, Aweme, Manitoba, USNM 17668; 21, E. xandana, New 
Brighton, Pennsylvania, CH 8 Oct. 1924 (USNM); 22, E. vertumnana, Merivale, Ontario, 
RLB slide 569 (CNC); 23, E. sotipena, Cincinnati, Ohio, RLB slide 575 (ANSP); 24, E. 
laracana, Dallas, Texas, RLB slide 573 (MCZ); 25, E. laracana, Cincinnati, Ohio, USNM 
slide 17719. 


338 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





FIGs. 26, 27. 
RLB slide 453 (ANSP); 27, E. bicordana, Aweme, Manitoba, RLB slide 552 (CNC). Scale 


lines = 1 mm. 


Female genitalia of Epinotia species. 26, E. zandana, Cincinnati, Ohio, 


Epinotia xandana; Heinrich (1923:204, as synonym of vertumnana), (1929:15). 
Eucosma yandana Kearfott (1907:25). 


Eucosma nothrodes Meyrick (1912:34, invalid replacement name). 
Epinotia yandana; Heinrich (1923:206, fig. 369), (1929:15, as synonym of xandana). 


Adult. 6 Types, xandana, yandana; 15 6, 18 ? examined (AB, ANSP, JRH, RLB, CNC, 
LACM, MEM, USNM). 


VOLUME 40, NUMBER 4 339 





Fics. 28, 29. Female genitalia of Epinotia species. 28, E. xandana, eighth segment 
and papillae anales, Cincinnati, Ohio, RLB slide 462 (CNC); 29, E. laracana, corpus 
bursae, Dallas, Texas, RLB slide 458 (MCZ). 


Forewing (Fig. 14). FWL: 6.0-8.0 mm; length of costal fold 0.33-0.38 FWL (5n); 
grayish white or dark grayish brown with brown subbasal and median fasciae, some 
specimens with scattered brown scales forming reticulations between fascia. 

Male genitalia (Fig. 21). Width of uncus at bifurcation 0.25-0.27 greatest width of 
juxta (2n); dorsal anellar plate with straight to slightly sinuate lateral margins, not ex- 
panded apically; aedeagus with eight cornuti (ln); valva abruptly narrowed beyond 
sacculus; saccular spine cluster elongate. Five preparations examined (ANSP, RLB, USNM). 

Female genitalia (Fig. 28). Tergum VIII sclerotized laterally, forming two narrow 
tergites, with irregular row of setae on posterior margins of tergites, anterior margin of 
each tergite straight, forming acute angle with anterior apophyses; papillae anales as in 
bicordana, except slightly less setose; length of anterior apophyses 0.88-0.97 length of 
posterior apophyses (4n); posterior margin of sterigma concave; seminalis bursae without 
spinules; width of posterior signum 2.0—2.2 width of anterior signum (3n). Six prepara- 
tions examined (ANSP, CNC, RLB, USNM). 

Types. Eucosma xandana—Lectotype, 4, genitalia CH 15 Dec. 1919. Type locality: 
Cincinnati, Ohio. Eucosma yandana—Lectotype, 4, genitalia slide “CH June 17, 1924”. 
Type locality: New Brighton, Pennsylvania. Both lectotypes designated by Klots (1942); 
in AMNH. 

Geographical distribution and flight times. Western Pennsylvania and Ohio (3 Mar.- 
12 Apr.) to N Mississippi (21-24 Feb.), and E Texas (27 Feb.). 

Host. Unknown. 


Discussion. The grayish white forewing was thought to distinguish 
this species from all others. Dark specimens recently collected with 
grayish white specimens in Mississippi are similar in fasciae, reticulated 
lines between fasciae, length of forewing costal fold, and in male and 
female genitalia. A uniformly dark female in poor condition is among 
the co-types of Zeller’s Paedisca vertumnana, and the partially dam- 
aged genitalia suggest it should be assigned to this species. These dark 
specimens indicate a degree of variation similar to that in related species; 


340 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


this species differs from others in having a light form with transverse 
fasciae rather than a longitudinal dark streak. The female is similar to 
E. zandana and E. bicordana in having papillae anales that are not 
reduced, and posterior and anterior apophyses that are subequal in 
length; it resembles E. vertumnana, E. laracana, and E. sotipena in 
having the tergum reduced to lateral tergites. The papillae anales of 
some females are covered with debris, similar to E. zandana. The male 
genitalia are similar to those of E. vertumnana, but differ in having a 
narrower uncus, a dorsal anellar plate not expanded apically, and a 
wider saccular spine cluster. 


Epinotia laracana (Kearfott) 
(Bigs 12, aye24 2529) 


Proteopteryx laracana Kearfott (1907:45). 

Proteopteryx navalis Meyrick (1912:34, invalid replacement name). 

Paedisca vertumnana Zeller (1875:310, part, var. e, f). 

Epinotia vertumnana; Heinrich (1923:204, part, ine. fig. 371). 

Epinotia atristriga; Clarke (1953:228, part, fig. 3b). 

Paedisca celtisana Riley (1881 [1882]:319); Fernald (1902 [1903]:459, as Eucosma); Hein- 
rich (1923:204, as synonym of vertumnana). REVISED SYNONYMY. 


Adult. 2 Type laracana, 6 type, celtisana; 22 6, 48 2? examined (AMNH, ANSP, CNC, 
ECK, INHS, MCZ, MEM, MSU, RLB, USNM). 

Forewing (Figs. 12, 13). FWL: 6.1-7.2 mm; length of costal fold 0.39-0.45 FWL 
(20n); light grayish brown to grayish brown intermixed with variable amounts of white 
or white-tipped scales; narrow to broad brown subbasal fascia extending from middle of 
discal cell to inner margin; narrow to broad brown median fascia usually extending from 
costa to CuA1, usually widened in discal cell; brown preapical and apical spots present 
or absent; most specimens with longitudinal streak between wing base and basal fascia, 
some specimens with sinuate, longitudinal streak between wing base and basal fascia, 
some specimens with sinuate, longitudinal streak extending from base through subbasal 
and median fasciae to preapical spot, some longitudinally streaked specimens with large, 
white spot at middle of inner margin (Fig. 13); some specimens suffused with grayish 
orange between median fascia and apex. 

Male genitalia (Figs. 24, 25). Width of uncus at bifurcation 0.26—0.35 width of juxta; 
dorsal anellar plate with sinuate lateral margins; aedeagus with 14-17 cornuti (13n); 
valva tapered or narrowed near middle, length of cucullus less than or subequal length 
of sacculus; saccular spine cluster elongate. Thirty-seven preparations examined (AMNH, 
CNC, ECK, MCZ, MSU, MEM, RLB, USNM). 

Female genitalia (Fig. 29). Tergum VIII as in xandana; papillae anales reduced, 
posterior half facing ventrally, anteriorly narrowed and facing laterally; length of ante- 
rior apophyses 0.70-0.79 length of posterior apophyses (28n); posterior margin of lamella 
postvaginalis V-shaped, acutely angled; bursa seminalis with sparse, small spinules; width 
of posterior signum 1.0-1.4 width of anterior signum (25n). Thirty-two preparations 
examined (AMNH, BM, CNC, ECK, MCZ, MSU, USNM). 

Types. Proteopteryx laracana—Lectotype, 2, designated by Klots (1942); in AMNH. 
Type locality: Cincinnati, Ohio. Paedisca celtisana—Holotype, 6, in USNM. Type lo- 
cality: Dallas, Texas. E. laracana was described from 13 specimens, 8 of which are E. 
vertumnana. 

Geographical distribution and flight times. Southern Ontario (26 Apr.) to E Pennsyl- 
vania (19 Apr.), W to Wisconsin (20 May), and S to central Mississippi (8 Mar.) and E 
Texas (10 Feb.). 

Host. Celtis (holotype of celtisana). 


VOLUME 40, NUMBER 4 341 


Discussion. Identities of E. laracana, E. vertumnana, and a new 
species, E. sotipena, have been confused frequently. Males of E. lar- 
acana can be identified by characters including forewing maculation 
(Figs. 12, 13), more tapered valva, aedeagus with 14-17 cornuti (more 
than any related species), and dorsal anellar plate with sinuate lateral 
margins. Females are distinctive among related species in having signa 
subequal in width. E. laracana appears to have a flight period concur- 
rent with E. sotipena, and slightly later in the spring than E. vertum- 
nana. 


Epinotia sotipena Brown, new species 


(Figs. 15, 23, 31) 


Proteopteryx laracana Kearfott (1907:45, part). 
Epinotia laracana; Heinrich (1923:204). 


Adult. 21 6, 45 ¢ examined (AMNH, ANSP, CNC, CU, INHS, JGF, JRH, RLB, USNM). 

Forewing (Fig. 15). FWL: 5.8-8.2 mm; length of male costal fold 0.31-0.85 FWL 
(15n); light grayish brown intermixed with variable amounts of white or white-tipped 
scales; brown basal streak extending from base to 0.39-0.45 FWL; curved, brown, median 
fascia extending from costa to CuA1; brown preapical spot present or absent, confluent 
with median fascia in some specimens; small, brown apical spot present or absent, con- 
fluent with preapica! spot in some specimens, strigulae on basal third and apical third of 
costa, basal row of outer marginal scales, and scattered scales in tornus dark grayish 
brown, most specimens suffused with grayish orange between median fascia and apex, 
some specimens also suffused with grayish orange between base and median fascia. 

Male genitalia (Fig. 23). Width of uncus at bifurcation 0.24—-0.30 greatest width of 
juxta; dorsal anellar plate and valva as in laracana; aedeagus with 12-14 cornuti; saccular 
spine cluster elongate. Nine preparations examined (ANSP, CNC, INHS, JGF, USNM). 

Female genitalia (Fig. 31). Tergum VIII as in E. xandana, except lateral tergites 
narrowed medially, anterior margin of each tergite forming obtuse angle with anterior 
apophyses; papillae anales as in E. laracana; length of anterior apophyses 0.60-0.69 
length of posterior apophyses; lamella postvaginalis as in E. laracana; bursa seminalis 
with dense, large spinules; width of posterior signum 2.0-3.3 width of anterior signum 
(5n). Nine preparations examined (CNC, INHS, USNM). 

Holotype. 2, Plummers Id. [Island], Md. [Montgomery Co., Maryland], 7-IV-62, R. W. 
Hodges. U.S.N.M. Type No. 76280; in USNM. Data given as on the label except for 
bracketed information. Specimen is not dissected, is in excellent condition, and has a 
forewing length of 7 mm. 

Paratypes. Arkansas: Johnson Co., 9 mi [14.5 km] N Clarksville, 11 Mar. 1985, R. L. 
Brown (1 é, genit. slide RLB 1425). Illinois: Putnam Co., 1 May 1987 (1 2, genit. slide 
USNM 17718), 2 Apr. 1989 (1 4, genit. slide RLB 446), 5 Apr. 1989 (1 4, genit. slide CH 
13 Aug. 1940, #8), 9 Apr. 1939 (1 9), 14 Apr. 1940 (1 4), 21 Apr. 1940 (1 8, genit. slide 
RLB 456), 28 Mar. 1948 (1 2, genit. slide RLB 166). Maryland: same data as Holotype 
(2 6, 2 8, 2 genit. slide USNM 17674). Missouri: Jackson Co., Independence, 12 Apr. 
1969, J. R. Heitzman (1 6), Kansas City, 13 Apr. 1969 (1 4); Jasper Co., Sarcoxie, city 
limits, open field, 15 Apr. 1975, R. Letsinger (1 2). New York: Tompkins Co., Ithaca, Six 
Mile Creek, 20 Apr. 1957, J. G. Franclemont (1 4, 1 9, 6 genit. slide JGF 4478), 22 Apr. 
1957 (1 6), 24 Apr. 1957 (1 4, 1 2), 26 Apr. 1957 (1 2), 25 Apr. 1959 (2 4, 3 Q, 6 genit. 
slide RLB 576), 16 Apr. 1960 (1 2), 7 May 1961 (1 2), same data except R. W. Hodges, 
18 Apr. 1959 (1 9), 27 Apr. 1961 (1 2, genit. slide USNM 17617), 7 May 1961 (1 4, 3 9), 
12 May 1961 (1 2), Buttermilk Falls, Apr. 1920 (1 6). Ohio: Hamilton Co, Cincinnati, 19 
Mar. 1908, Annette F. Braun (2 2, genit. slide CNC Epi. 12), 26 Mar. 1903 (1 @), 15 Apr. 
1904 (1 2), 17 Mar. 1905 (1 4, genit. slide CNC 12), 8 Apr. 1906 (2 6, 9 2), 20 Mar. 1904 


342 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 30, 31. Female genitalia of Epinotia species. 30, E. vertumnana, St. Davids, 


Ontario, RLB slide 460 (CNC); 31, E. sotipena, paratype, Putnam Co., Illinois, USNM 
slide 17718. 


(1 4, genit. slide RLB 575); Warren Co., 15 Mar. 1935, Annette F. Braun (1 2). Penn- 
sylvania: Beaver Co., New Brighton, 12 Apr. 1902, H. D. Merrick (1 2), 15 Apr. 1902 (2 
2), 3 Apr. 1908 (1 4, genit. slide USNM 17675), 10 May 1907 (1 2); Dauphin Co., Rockville 
(1 4, 1 2, 6 genit. slide CH 29 Oct. 1921, #2). Ontario: Ottawa East, 29 Apr. 1944, J. 
McDunnough (1 2, genit. slide RLB 447). Quebec: Chelsea, 24 Apr. 1933, G. S. Walley 
(1 2), 21 Apr. 1933, J. McDunnough (1 2, genit. slide RLB 450), Old Chelsea, 3 May 
1939, T. N. Freeman (1 2). Paratypes are deposited in collections listed in the material 
examined for the adult, and in BMNH. 
Host. Unknown. 


VOLUME 40, NUMBER 4 343 


Discussion. Male genitalia of this superficially distinctive species are 
similar to those of E. laracana. Both species have valvae that intergrade 
in form from gradually narrowed to somewhat abruptly narrowed, 
although the abruptness of narrowing is less in both than in E. vertum- 
nana. Width of uncus relative to juxta is usually greater in E. laracana 
than in E. sotipena. The aedeagus has 14-17 cornuti in E. laracana 
and 12-14 in E. sotipena. Males of the two species are easily separated 
by length of the costal fold relative to forewing length, the fold being 
shorter in E. sotipena. Female E. sotipena differ from E. laracana in 
having anterior apophyses shorter relative to posterior apophyses, bursa 
seminalis with large spinules, and posterior signum much wider than 
anterior signum. Some females have debris on the papillae anales, as 
in E. zandana. 


Epinotia vertumnana (Zeller) 
(Figs. 8, 16, 17, 22, 30) 


Paedisca vertumnana Zeller (1875:310, var. d). 

Epinotia zandana; Heinrich (1923:205, fig. 370); Comeau & Roelofs (1973:197); Roelofs 
& Cardé (1974:98) (not Kearfott). 

Epinotia prob. zandana; Comeau & Roelofs (1978:194). 

Epinotia atristriga Clarke (1953:228, part). 

Epinotia atistriga; Comeau & Roelofs (1973:194); Roelofs & Cardé (1974:98) (misspelling 
and misidentification). 

Epinotia atristriga; Roelofs & Brown, 1982:411 (not Clarke, 1953). 

Epinotia sp.; MacKay (1959:64). 


Adult. 2 Type; 148 6, 117 2 examined (AMNH, ANSP, CNC, CU, INHS, MEM, MSU, 
RLB, UCB, USNM). 

Forewing (Figs. 16, 17). FWL (nonreared): 5.5-7.0 mm; length of costal fold 0.36- 
0.42 FWL (22n); uniformly grayish brown or grayish brown peppered with varying 
numbers of white-tipped scales, some specimens with discontinuous, dark grayish brown, 
basal, median, and apical! longitudinal streaks or with continuous longitudinal streak; 
rarely with dark grayish brown, subbasal and median fasciae, and preapical spot. 

Male genitalia (Figs. 8, 22). Width of uncus at bifurcation 0.21—0.24 width of juxta 
(15n); dorsal anellar plate with straight lateral margins, apically curved outwardly; ae- 
deagus with 6-10 cornuti (22n); valva abruptly narrowed beyond sacculus, cucullus dis- 
tinctly longer than sacculus; saccular spine cluster elongate. Forty-seven preparations 
examined (CNC, MSU, RLB, USNM). 

Female genitalia (Fig. 30). Tergum VIII as in E. xandana; papillae anales as in E. 
laracana; \ength of anterior apophyses 0.67-0.77 length of posterior apophyses (15n); 
posterior margin of sterigma irregularly concave; bursa seminalis without spinules; width 
of posterior signum 1.7-2.5 width of anterior signum (14n). Twenty-two preparations 
examined (AMNH, CNC, CU, RLB, USNM). 

Larva. The description of Epinotia sp. by MacKay (1959) was based on nine larvae 
reared from Crataegus at St. David’s and Merivale, Ontario. Associated adults from both 
localities are identified here as E. vertumnana. 

Type. Lectotype here designated: 2, Dallas, Tex., Boll; vertumnana var. d [green, 
handwritten label]; genitalia slide R. L. Brown 1158 [green label], Type 14336 [red label], 
Lectotype Paedisca vertumnana by R. L. Brown [red bordered label]; in MCZ. The 
original corroded pin holding the specimen has been clipped and the specimen has been 
double-mounted on a polyporus block. The specimen is in good condition, except that 
the right wings are rubbed. The lectotype is similar to the specimen in Fig. 16, except 
the median and apical streaks are continuous, and the basal streak is narrower. 


344 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Geographical distribution and flight times. Southern Ontario and Quebec (19 Mar.— 
20 Apr.) to E Pennsylvania (14 Mar.-15 Apr.), W to Michigan (23-31 Mar.), and S to N 
Mississippi (21-24 Feb.) and E Texas. Two males (UCB) tentatively identified as this 
species were collected during early March in Jefferson Co., Colorado. 

Host. Crataegus (72n), reared by P. J. Chapman and S. E. Lienk in New York, and 
in Ontario by W. L. Putnam, T. N. Freeman, and J. McDunnough. 


_Diseussion. Zeller described six varieties (a-f) of this species from 
an unknown number of specimens collected in Texas by Jacob Boll and 
one specimen collected by Speyer in New York. The type series in MCZ 
includes two specimens of “var. a’’, one specimen of “‘var. c’’, one 
specimen of “var. d’’, five specimens of “var. e’’, and two specimens 
of ‘“‘var. f’, all from Dallas, Texas. Specimens of “var. e’”’ and “var. f” 
are conspecific with E. laracana, and “‘var. c’’ is identified tentatively 
as E. xandana. The longitudinally streaked “var. d’”’ specimen, desig- 
nated as lectotype, appears clearly conspecific with specimens reared 
from Crataegus. Specimens of “var. a” tentatively are considered syn- 
onymous with the lectotype, but these males differ in having a con- 
trastingly paler inner margin of the forewing. 

The unicolorous and fasciate forewing forms of E. vertumnana were 
misidentified by Heinrich (1928) and others as E. zandana. Specimens 
with longitudinally streaked forewings were named by Clarke (1953) 
as E. atristriga, a junior synonym of E. zandana. Two male specimens 
from Colorado have a superficially darker appearance, similar to E. 
zandana, but their genitalia cannot be differentiated from other E. 
vertumnana; collection of females from this area should clarify their 
identity. 

In addition to smaller size and lighter color, E. vertumnana differs 
from E. zandana in the male genitalia by having a narrow uncus and 
an elongate saccular spine cluster, and, in the female, by having re- 
duced papillae anales and long anterior and posterior apophyses. E. 
vertumnana differs from E. laracana and E. sotipena in having less 
than 11 cornuti and a dorsal anellar plate with straight lateral margins 
in the male. Females of E. vertumnana differ from those of E. laracana 
in having one signum much wider than the other, and from those of 
E. laracana and E. sotipena in having the posterior margin of the 
lamella postvaginalis irregularly concave rather than acutely angled. 

Although the pheromone of E. vertumnana has not been identified, 
males are attracted to cis-7-dodecenyl acetate. Because this species was 
misidentified as “Epinotia sp. prob. zandana” and “E. atistriga’’, the 
attractancy of males to another form of females was considered to be 
an example of cross-attractancy between species (Comeau & Roelofs 
1973). Fortunately, voucher specimens of both forms were retained 
(CU) and the correct identity now can be established. 


VOLUME 40, NUMBER 4 345 


ACKNOWLEDGMENTS 


I thank Frederick H. Rindge (AMNH), Donald Azuma (ANSP), Bryant Mather, L. L. 
Pechuman (CU), John G. Franclemont, J. Richard Heitzman, George L. Godfrey (INHS), 
Edward C. Knudson, Julian P. Donahue (LACM), Margaret K. Thayer (MCZ), Frederick 
W. Stehr (MSU), Jerry A. Powell (UCB), and Donald R. Davis (USNM) for the loan of 
specimens. The assistance of Gaden Robinson and Kevin Tuck at the British Museum 
(Natural History) is appreciated. Photographs of imagos were made by Howard Lyons; 
drawings by David Adamski (Fig. 8) and Amy Trabka. This research was supported in 
part by National Science Foundation Grant DEB 77-15808 and a Grant-in-Aid of Re- 
search from Sigma Xi, the Scientific Research Society of North America. 


LITERATURE CITED 


BARNES, W. & J. MCDUNNOUGH. 1917. Check list of the Lepidoptera of boreal Amer- 
ica. Herald Press, Decatur, Illinois. viii + 392 pp. 

Busck, A. 1902 [1903]. In Dyar, H. G., A list of North American Lepidoptera. U.S. 
Natl. Mus. Bull. 52:xix, 1-723. 

1903. A revision of the American moths of the family Gelechiidae, with de- 
scriptions of new species. Proc. U.S. Natl. Mus. 25:767-930. 

CLARKE, J. F. G. 1953. New species of Olethreutidae from Illinois (Lepidoptera). J. 
Wash. Acad. Sci. 43:226-281. 

CLEMENS, B. 1861 [1862]. Micro-lepidopterous larvae. Proc. Entomol. Soc. Phila. 1: 
5-87. 

1864 [1865]. North American micro-lepidoptera. Proc. Entomol. Soc. Phila. 3: 
505-520. 

ComMEAuU, A. & W. L. ROELOFS. 1978. Sex attraction specificity in the Tortricidae. 
Entomol. Exp. Appl. 16:191—200. 

CoTk, W. A. & D.C. ALLEN. 1973. Biology of the maple trumpet skeletonizer, Epinotia 
aceriella (Lepidoptera: Olethreutidae), in New York. Can. Entomol. 105:463—470. 

FELT, E. P. 1905. Insects affecting park and woodland trees. N.Y. State Mus. Mem. 8, 
vol. 1:1—459. 

FERNALD, C. H. 1882. A synonymical catalogue of the described Tortricidae of North 
America, north of Mexico. Trans. Am. Entomol. Soc. 10:1-72. 

1902 [1903]. In Dyar, H. G., A list of North American Lepidoptera. U.S. Natl. 

Mus. Bull. 52:xix, 1-723. 

1908. The genera of the Tortricidae and their types. Carpenter & Morehouse, 
Amherst, Massachusetts. 68 pp. 

HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucos- 
minae of the family Olethreutidae. U.S. Natl. Mus. Bull. 123:iv, 1-298. 

1929. Notes on some North American moths of the subfamily Eucosminae. 
Proc. U.S. Natl. Mus. 75:1-28. 

KEARFOTT, W. D. 1907. New North American Tortricidae. Trans. Am. Entomol. Soc. 
33:1-98. 

Kuiots, A. B. 1942. Type material of North American microlepidoptera other than 
Aegeriidae in the American Museum of Natural History. Bull. Am. Mus. Nat. Hist. 
79:391-424. 

JOHNSON, W. T. & H. H. Lyon. 1976. Insects that feed on trees and shrubs. Cornell 
Univ. Press, Ithaca, New York. 464 pp. 

MacKay, M. R. 1959. Larvae of the North American Olethreutidae (Lepidoptera). 
Can. Entomol. Suppl. 10:1-338. 

1962. Additional larvae of the North American Olethreutinae (1) (Lepidoptera: 
Tortricidae). Can. Entomol. 94:626-643. 

MEYRICK, E. 1912. On some impossible specific names in micro-lepidoptera. Entomol. 
Monthl. Mag. ser. 2, 48:32-36. 




















346 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


MILLER, W. E. 1973. Clemens types of Olethreutinae (Lepidoptera, Tortricidae). Trans. 
Am. Entomol. Soc. 99:205-234. 

Oxu, T. 1979. Acroclita and some allied genera (Lepidoptera: Tortricidae) from Japan, 
with descriptions of new taxa. Kontyi 47:586-592. 

RILEY, C. V. 1881 (1882). Descriptions of some new Tortricidae (leaf rollers). Trans. 
St. Louis Acad. Sci. 4:316-324. 

ROELOFS, W. L. & R. L. BROWN. 1982. Pheromones and evolutionary relationships of 

_ Tortricidae. Ann. Rev. Ecol. Syst. 18:395-422. 

ROELOFS, W. L. & R. T. CARDE. 1974. Sex pheromones in the reproductive isolation 
of lepidopterous species, pp. 96-104. In Birch, M. C. (ed.), Pheromones. American 
Elsevier, New York. 

ZELLER, P. C. 1875. Beitrage zur Kenntniss der nordamericanischen Nachtfalter be- 
sonders der Microlepidopteren. Verh. Zool.-bot. Ges. Wien 25:207-360, pls. 8-10. 


Received for publication 4 August 1986; accepted 10 October 1986. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
40(4), 1986, 347 


MYNES GEOFFROYI GUERINI WALLACE (NYMPHALIDAE) 
PARASITIZED BY A TACHINID FLY 


Mynes geoffroyi guerini Wallace, the white nymph, is a poorly studied nymphalid 
that occurs in coastal rainforests sporadically from the Claudie River in northern Queens- 
land to near Ballina, northern New South Wales (Common & Waterhouse 1981, Austra- 
lian butterflies, Angus & Robertson, 651 pp.). The larvae are gregarious and feed exclu- 
sively on the young foliage of the tall stinging trees Dendrocnide moroides (Wedd.) 
Chew and D. photinophylla (Kunth) Chew, and the nonstinging rainforest shrub Pip- 
turus argenteus (Forster) Wedd. (Common & Waterhouse, cited above). There are four 
Australian species of Dendrocnide, while Pipturus is only represented by one species; 
both genera are from the stinging nettle family Urticaceae. 

On 25 May 1985 I collected one larva of Mynes geoffroyi guerini from a young leaf 
of a mature P. argenteus in disturbed rainforest near Mt. Coot-tha, about 5 km W of 
Brisbane, Queensland. The larva, a last instar, had a fly egg behind the head. Another 
nine larvae were observed, but close examination showed they had not been parasitized. 
The parasitized larva as well as three unparasitized ones were taken to the laboratory 
and placed in a plastic bag with fresh leaves of the food plant. The parasitized larva 
pupated on 27 May. Two days later, it displayed no movement when handled (normal, 
healthy pupae of M. g. guerini usually thrash their bodies wildly about for up to 20 
seconds if disturbed) and was subsequently broken to disclose an active fly maggot, which 
pupated 1 June 1985. Length of the fly pupa was 6.8 mm. The other butterfly larvae 
were reared successfully to the adult stage. The fly emerged on 6 July, after 35 days in 
the pupal stage. The fly was deposited in the collection of the Entomology Department, 
Department of Primary Industries (DPI), Indooroopilly, Queensland, and was identified 
by Dr. B. K. Cantrell of that Department as Compsilura concinnata (Meigen) (Tachin- 
idae). 

This appears to be the first published record of fly parasitism of M. geoffroyi guerini. 
Compsilura concinnata has been recorded as a parasite of the following Australian moths 
by Crosskey (1978, Bull. Brit. Mus. Nat. Hist. (Entomol.) Suppl. 21:1-221)—Doratifera 
vulnerans Lewin (Limacodidae) and Anomis xanthindyma Boisduval and Brithys crini 
(Fabricius) (both Noctuidae). Recently, Chadwick and Nikitin (1985, Aust. Zool. 21:587- 
598) recorded the Australian moth Isotenes miserana (Walker) (Tortricidae) and the 
introduced butterfly Artogeia (Pieris) rapae (L.) (Pieridae) as pupal hosts of C. concin- 
nata. 

The parasitoid fly C. concinnata has a widespread distribution, and is known from 
the Palearctic region (including Japan), Africa, the Oriental Region to Australia, and has 
been introduced into North America (Arnaud 1969, Pan-Pacific Entomol. 45:77; 1978, 
USDA Misc. Pub. No. 1319:149-168). The species is one of the most widely reared 
parasites of Lepidoptera in Europe (Herting 1960, Mono. Ang. Entomol. Nr. 16:55-57) 
and North America (Arnaud, cited above). These authors each record over 100 species 
of lepidopteran hosts from a wide range of families and genera of butterflies and moths, 
indicating that C. concinnata is a generalist and opportunistic fly in larval feeding habits 
and the type of habitat frequented. Although Australian host records are scanty, they do 
show similar trends of nonspecificity to any one group of Lepidoptera. 

The pupal duration of 35 days of C. concinnata recorded here is considerably longer 
than the 11-14 days recorded for another tachinid fly parasite of Australian butterflies, 
Winthemia neowinthemioides (Townsend) (Smithers 1973, Aust. Entomol. Mag. 1:37- 
40; Hawkeswood 1986, in review). Whether this longer pupation period results from 
species-specific differences or is due to the colder conditions under which C. concinnata 
was reared (during winter) awaits further research. 


T. J. HAWKESWooD, 49 Venner Road, Annerley, Brisbane, Queensland, Australia, 
4108. 


348 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Journal of the Lepidopterists’ Society 
40(4), 1986, 348-349 


AN OVIPOSITION “MISTAKE” BY BATTUS PHILENOR L. (PAPILIONIDAE) 


Although oviposition on plants unsuitable for larval development occurs among the 
Lepidoptera (Berenbaum 1981, J. Lepid. Soc. 35:75 and references therein), the actual 
egg-laying episodes have rarely been witnessed. I report here direct field observation of 
an oviposition “mistake” by the pipevine swallowtail, Battus philenor. This record is 
particularly noteworthy because the errant female was one for which a detailed field 
record of egg-laying activity over three weeks was accumulated. 

On 7 April 1981, at a field site in Kirby State Forest, Tyler Co., Texas, Mark D. 
Rausher and I observed a Battus female deposit a single egg on a leaf of Smilax laurifolia 
L. (Liliaceae). This event represents the only oviposition by this butterfly species on a 
plant outside the genus Aristolochia (Aristolochiaceae) noted in more than eight years 
of fieldwork on the Texas population by me and colleagues. Our studies included obser- 
vations of hundreds of ovipositing butterflies and tens of thousands of landings on nonhost 
foliage. No Aristolochia plants were present within 2 m and the female had not discov- 
ered any host plants in the previous 2 min of observation. Although I intended to follow 
the progress of larval development on the plant, both egg and leaf were gone on the 
third day after oviposition. Nevertheless, 25 larvae freshly hatched from eggs laid by 
captive females on the native E Texas host plants, A. reticulata Nutt. and A. serpentaria 
L., and placed on cuttings of young Smilax foliage failed to consume any leaf material 
and did not survive to the second instar. By contrast, another 25 larvae fed on cuttings 
of both Aristolochia hosts progressed normally through the first instar and many even- 
tually pupated. 

Most remarkable about the egg deposition on Smilax was the insect’s extended period 
of investigation; several minutes of fluttering over the leaf and circling about the plant 
preceded the aberrant oviposition. During oviposition, Battus females alight frequently 
on nonhost plants upon which they drum their foretarsi, presumably tasting the leaf 
surface with tarsal chemoreceptors (Feeny, Rosenberry & Carter 1983, pp. 27-76 in 
Herbivorous insects: Host-seeking behavior and mechanisms, Academic Press, New York, 
and references therein). Most individuals leave nonhost plants immediately after landing 
and resume searching for host plants. 

The extensive history of searching by this particular Battus female suggests several 
explanations for the anomalous egg deposition. First, the female may have been more 
prone to oviposit on an unsuitable plant due to low rates of discovery of the principal 
Aristolochia hosts. The rate of discovery of Aristolochia plants in the minutes before the 
Smilax oviposition was indeed low (0.19 host plants/min vs. a mean of 1.02 host plants/ 
min for 6 other observation periods). Since no eggs were laid on several hosts discovered 
just minutes earlier, however, the female was not evidently “desperate” to lay eggs. 

Alternatively, the female may have suffered some deterioration of the sensory appa- 
ratus required to identify a host plant after landing. Extremely worn and tattered at the 
time she oviposited on Smilax, the female had been first marked two weeks previously. 
Only 1 other butterfly of almost 200 marked was followed as long. The efficiency with 
which the aging female landed on host plants did indeed decline in successive recaptures 
(Fig. 1). Since relative host density did not change significantly over this period, the 
female was apparently identifying host plants before landing less accurately as she aged. 
ete she was discriminating between hosts and nonhosts less accurately after landing 
as well. 

Finally, the mistaken deposition may have resulted from the female’s previous expe- 
rience with host plants. Pipevine swallowtail butterflies learn to search for the leaf shape 
of preferred host plants (Papaj 1986, Anim. Behav. 34:1281-1288). This particular fe- 
male, for example, almost always alighted on broad nonhost leaves resembling the broad- 
leaved A. reticulata (Fig. 1). The Smilax plant on which an egg was laid consisted of a 
solitary, newly formed broad leaf on a short stem, and bore a striking visual resemblance 


VOLUME 40, NUMBER 4 349 


Proportion Proportion 
1 142 10 39 3! 
1.0 a C= 6 555-455 CS @ ----------- P) 1.0 
r te bigoe: leaves 
8 e---e e alighted on 5 


Host plants 
ye alighted on 





20 25 30 5 | lo 15 
MARCH APRIL 


Day Followed 


Fic. 1. Record of host-landing efficiency and leaf-shape response over time. Solid 
line indicates time course of host-landing efficiency, expressed as proportion of all plants 
alighted on that were Aristolochia. Dotted line indicates time course of leaf-shape pref- 
erence, expressed as fraction of all nonhost leaves alighted on that were broad. Number 
above each point indicates sample size on each observation day. Arrow indicates day of 


anomalous egg deposition. 


in color, leaf shape, and growth form to phenologically young A. reticulata plants which 
are highly preferred for oviposition (Rausher 1980, Evolution 34:342-355; Papaj & Raush- 
er 1987, Ecology, in press). Possibly, the experienced female became so positively re- 
sponsive to the visual stimuli of young A. reticulata foliage (which the young Smilax 
leaf mimicked well enough to deceive two human observers) that conflicting negative 
responses to nonhost chemotactile cues were eventually suppressed. 


DANIEL R. PAPaj, Department of Zoology, Duke University, Durham, North Carolina 
27706. CURRENT ADDRESS: Department of Entomology, Fernald Hall, University of 


Massachusetts, Amherst, Massachusetts 01003. 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Journal of the Lepidopterists’ Society 
40(4), 1986, 350-351 


EMERGENCE OF TISCHERIA IMMACULATA (BRAUN) (TISCHERIIDAE) 
FROM LEAVES OF CEANOTHUS GRISEUS L. 


-Tischeria immaculata (Braun) larvae mine leaves of Ceanothus griseus L., a native 
ornamental in Washington State and California (Braun 1915, Entomol. News 26:271- 
273; Fasoranti 1983, Z. Angew. Entomol. 96:270-476). Biology and mining habits were 





Fics. 1-6. Emergence of Tischeria immaculata. 1, 2, Insect thrusts itself from the 
mine and withdraws legs; 3, Insect falls on back and rests for a few seconds; 4, It flips 
over suddenly and stays in this position for about 15 seconds; 5, Wing pumping, expan- 
sion, and drying accompanied by circular movements for 45 seconds; 6, Wing beating 
before insect rests. 


VOLUME 40, NUMBER 4 S15) 


elucidated in Washington by Fasoranti (1984, Can. Entomol. 116:1441-1448). Adults 
appear at the beginning of May, and there is a short cycle of development during the 
warm weather between May and August. The next generation takes longer, from August 
through May of the following year. Mating and egg laying starts immediately after 
emergence. 

Emergence takes place mostly in the evening (1750-2400 h). Time of emergence 
provides some protection against the insect’s principal predator, the dark eyed junco, 
Junco hyemalis L. Emergence also is temperature dependent. In field and laboratory, 
emergence occurred only at temperatures between 20-22°C. Insects rarely emerged be- 
low or above these temperature limits. A typical exit hole is crescentic, about 0.15-—0.20 
mm long (N = 350), with the convex side toward the end of the mine. The center of the 
crescent is about 0.20 mm from the end of the mine. 

Under controlled conditions (21°C and 70% RH), emergence of 75 adults was timed 
and photographically documented with a 35 mm camera attached to a microscope using 
Panatomic X film. The subjects were illuminated with light from two opposing sources. 
A typical sequence of emergence is shown in Figs. 1 to 6. The whole process takes 
between 6-7 minutes. 


J. OLANIYAN FASORANTI, Department of Biological Sciences, University of Ilorin, P. 
M. B. 1515, Ilorin, Nigeria. 


Received for publication 1 February 1986; accepted 14 July 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 351-352 


UNUSUAL PREDATOR DAMAGE TO CARTEROCEPHALUS SILVICOLUS 
(MEIGEN) (HESPERIIDAE) 


Bird-inflicted wing damage in Lepidoptera that fold their wings above their bodies at 
rest was classified into two categories by Beck and Garnett (1983, J. Lepid. Soc. 37:289- 
300). To continue the earlier classification proposed by Sargent (1976, Legion of night, 
Univ. Mass. Press, Amherst, Massachusetts, 222 pp.), who worked with noctuid species, 
the new categories were called Type IV (for bilaterally symmetrical tears roughly parallel 
to the main wing veins) and Type V (for beak imprints which cross the main wing veins 





Fic. 1. Carterocephalus silvicolus (Meigen) 4 (dorsal), showing Type Vb wing dam- 
age. 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Position of specimen when held by bird, reconstructed from cut-out tracings 
of the four wings. 


at roughly right angles). A subcategory of Type V (Type Vb) was proposed to accom- 
modate beak imprints across all four wings. We assumed this situation would always 
result in a successful capture, and Type Vb was included only for completeness. 

Since then, a specimen showing “hypothetical” Type Vb damage came to our attention 
(Fig. 1). Unfortunately, nothing is known about the circumstances surrounding its capture 
other than the data on the specimen label: Akademici Raros, Novaibirsok, USSR; 20-vi- 
1978; coll; V. Dubatolov. Despite this, two facts are clear. First, the insect was grasped 
by a bird’s beak firmly enough to leave distinct areas devoid of scales on all four wings. 
Second, the insect was subsequently released. We believe release was not caused by the 
insect’s efforts to escape, since no significant tears or other signs of pulling free are 
discernible. It is also unlikely that the insect startled the bird since all wings were im- 
mobilized and, from the position of the beak imprints, the legs and antennae were 
directed away from the bird’s face (Fig. 2 or its mirror image). 

Hypotheses explaining the skipper’s release include 1) an error in prey manipulation 
by the bird, 2) the possibility of some allomonal product of the skipper, and 3) an extrinsic 
startle from outside the predator-prey system. The first is possible and the second un- 
likely, since we know of no references to vertebrate-effective allomonal products pro- 
duced by members of this genus. We prefer the third possibility. If the bird was not 
processing prey at the site of capture but bringing it instead to nestlings, there would be 
time during which the bird could have been startled into releasing its prey by some 
extrinsic factor. 

We thank Balint Zsolt of Budapest, Hungary, for the C. silvicolus specimen, currently 
in the collection of the senior author, and ultimately to be deposited in the Florida State 
Collection of Arthropods, Gainesville, Florida, USA. 


ANDREW F. BECK, United States Navy Disease Vector Ecology and Control Center, 
Box 43, Naval Air Station, Jacksonville, Florida 322212-0043, AND WILLIAM J. GARNETT, 
2919 Cathedral Avenue NW, Washington, D.C. 20008. 


Received for publication 25 April 1986; accepted 29 July 1986. 


VOLUME 40, NUMBER 4 OD 


Journal of the Lepidopterists’ Society 
40(4), 1986, 353-354 


FOUR NEW UNITED STATES RECORDS OF MOTHS FROM TEXAS 


The four moths illustrated here represent new records for the United States and were 
collected in extreme S Texas in UV light traps. The lower Rio Grande valley has been 
drastically modified by the environmental changes that accompany rapid population 
growth, intensive agricultural land use, and widespread aerial spraying of pesticides. Few 
natural areas remain, and the consequences of this are becoming apparent. One endemic 
moth, Agapema solita Ferguson (Saturniidae), formerly common around Brownsville 
and known only from southern Texas (Ferguson in Dominick et al. 1972, Moths of 
America north of Mexico, fasc. 20.2B, E. W. Classey Ltd., London), has not been seen 
since 1956, and may now be extinct, or nearly so, as a result of habitat destruction. 
Probably there are other not as well documented examples. 

In spite of the loss of habitat, unusual species of Lepidoptera continue to appear, 
including many new records for the United States (Kendall & McGuire 1984, Bull. Allyn 
Mus. Entomol. 86:1-50; Blanchard & Knudson 1985, J. Lepid. Soc. 39:1-8). Some are 
probably individual nonbreeding immigrants from a large reservoir of species that still 
exists in the Sierra Madre Oriental of northeastern Mexico, where rich tropical to mon- 
tane temperate zone forests are found within 322 km of the border. However, some 
Mexican species do become established periodically in S Texas, later to be extirpated by 
low temperatures or drought. One case of temporary occurrence was reported but sub- 
sequently overlooked, and is perhaps worth citing here. Adults of the pantropical bean 
pod borer, Maruca testulalis (Geyer) (Pyralidae, Pyraustinae), were reared from larvae 
feeding on string beans at Olmito, Cameron Co., Texas (Williamson 1948, J. Econ. 
Entomol. 36:936—-937). This species has not again been reported from the continental 
United States, although it can thrive in agricultural environments in warmer climates 
(Ferguson 1988, Pests not known to occur in the United States or of limited distribution, 
No. 40, PPQ, Animal & Plant Health Inspection Service, USDA). Details of the new 
records follow. 

Euprosterna lacipea Druce (Limacodidae), Brownsville, Cameron Co.,7 August 1976, 
1 male; Santa Ana Nat'l. Wildlife Refuge, Hidalgo Co., 28 May 1982, 8 males (Fig. 1); 
14 May 1988, 1 male; 4 August 1986, 3 males; collected by E. C. Knudson. Body and 
wings dark brown, lightly frosted with light gray; bands on forewing upper side light 
gray; length of forewing 9-10 mm. This species appears well established in S Texas, 
though apparently was not present before 1976. There are three Mexican specimens in 
the U.S. National Museum (USNM) from Vera Cruz and Colima. 

Molybdogompha polymygmata Dyar (Geometridae), Santa Ana Nat'l. Wildlife Ref- 
uge, Hidalgo Co., 28 September 1980, 1 female (Fig. 2), collected by Knudson. Wing 
upper sides light yellowish brown with multiple, transverse parallel, blackish striations 
and a regular subterminal row of ocellate spots; terminal line of silvery, interrupted 
dashes; length of forewing 8.5 mm. The type specimen and a second specimen are in 
the USNM and both are from Vera Cruz. A third Mexican specimen was collected by 
Knudson near Cd. Valles, San Luis Potosi, 28 November 1978, and is in the American 
Museum of Natural History. The relations of this genus within the Geometridae are 
unclear, but its superficial appearance indicates that it might belong to the tribe Baptini 
(Ennominae). 

Eubaphe medea (Druce) (Geometridae), Bentsen State Park, Hidalgo Co., 27 May 
1982, 3 males (Fig. 3), collected by Knudson. Body orange; wings translucent pale orange 
with opaque bright orange discal patch; middle of forewing costal margin with fold; 
length of forewing 14-15 mm. This species has apparently been collected in Mexico, but 
the locality and present specimen location are not known to us. It is otherwise known 
from Guatemala, Honduras, Costa Rica, and Panama (Fletcher 1954, Zoologica [New 
York Zool. Soc.] 39:153-166). 

Psamathia placidaria (Walker) (Epiplemidae), Santa Ana Nat’l. Wildlife Refuge, 30 


354 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-4. 1, Euprosterna lacipea Druce; 2, Molybdogompha polymygmata Dyar; 
3, Eubaphe medea (Druce); 4, Psamathia placidaria (Walker). 


November 1981, 1 male (Fig. 4), collected by Knudson. Body and wings grayish brown 
above and below; wings above with innumerable, broken, blackish striations and black 
dots over lower median area of forewing and near anal margin of hindwing; hindwing 
produced as a short, black-marked, obliquely truncated tail; length of forewing 18 mm. 
This species ranges from Mexico to Venezuela. In Mexico, it has been collected in Ta- 
maulipas by Knudson and Alma Solis. 


DoucLas C. FERGUSON, Systematic Entomology Lab., USDA, % U.S. National Mu- 
seum of Natural History, Washington, D.C. 20560, AND EDWARD C. KNUDSON, 808 
Woodstock, Bellaire, Texas 77401, Research Associate, Florida State Collection of Ar- 
thropods, Gainesville, Florida 32602. 


Received for publication 12 May 1986; accepted 19 September 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 354-355 


NEW HOST RECORDS FOR EUPLOEA CORE CORINNA 
(MACLEAY) (NYMPHALIDAE) 


During the course of taxonomic studies on Asclepiadaceae and Apocynaceae, numerous 
observations of oviposition, feeding and pupation of the crow butterfly, Euploea core 
corinna (Macleay) were made. This butterfly is widely distributed in northern and east- 
ern Australia (Common & Waterhouse 1981, Butterflies of Australia, Angus & Robertson, 
Sydney) with temperature requirements and suitable host plants restricting the subspecies 
range (Scheermeyer 1985, Aust. J. Zool. 33:339-348). In a review of host plant records 
for this Australian species, various species of Asclepiadaceae and Apocynaceae, either 
naturalized or cultivated, were listed, as well as native species of these families and the 
Moraceae (Scheermeyer & Zalucki 1985, Aust. Entomol. Mag. 11:87—-90). Some host 
plants have been demonstrated to significantly affect development times, weights, size, 
and mortalities at all stages of the life cycle (Rahman, Zalucki & Scheermeyer 1985, J. 
Aust. Entomol. Soc. 24:95-98). 

Euploea core corinna is known at times to oviposit on unsuitable hosts (Kitching & 
Zalucki 1983, Aust. Entomol. Mag. 10:64—-66) and I have observed eggs on Mammillaria 
gracilis Pfeiff. (Cactaceae), but no larval feeding. In some instances, limited larval feed- 
ing may occur as observed on the Madagascan Cynanchum compactum Choux and three 


VOLUME 40, NUMBER 4 BaD, 


unidentified Cynanchum species related to C. mahafalense Jum. & Perr. (Asclepiada- 
ceae). 

I observed completion of the life cycle of the butterfly to the adult stage (new records 
indicated *) on the following hosts in cultivation at the localities and dates listed. 

1. In April 1986, on the African Adenium obesum Balf.* and A. multiflorum KI.* 
(Apocynaceae) at the Mt. Coot-tha Botanic Gardens, Brisbane, Queensland (27°29’S, 
153°00’E). Both young and mature leaves were eaten. No feeding was observed on 
adjacent plants of the related Madagascan Pachypodium lamieri Drake. 

2. During summer 1985-86 at Didcot, Queensland (25°28’S, 151°52’E), on the Austra- 
lian Sarcostemma australe R. Br. subsp. australe, S. australe tentative subsp. nov. 1 and 
S. australe tentative subsp. nov. 2 (Forster, unpubl.), but not on adjacent plants of the 
African S. viminale (L.) R. Br., S. vanlessenii Lavr., S. stolonifera Adams & Holland 
and S. socotranum Lavr. (Asclepiadaceae). Only young shoots were eaten. 

3. During summer 1985-86 at Annerley (27°31'S, 153°03’E) and Strathpine (27°24'S, 
152°57’E), Brisbane and Didcot, on young shoots and leaves of the Australian Hoya 
australis R. Br. ex Traill., H. sana F. M. Bail.* and H. macgillivrayi F. M. Bail.*, the 
New Guinean H. archboldiana C. Norman* and Asian H. carnosa (L.) R. Br.*, but not 
on adjacent plants of the Australian H. nicholsoniae F. Muell., H. poolei C. T. White & 
Francis and various unidentified species of Section Eriostemma (Asclepiadaceae). 

4. During late summer 1986 at Didcot on old leaves and shoots of Brachystelma 
microstemma Schltr.* (syn. Microstemma tuberosum R. Br., Forster 1985, Taxon 34: 
318-319) (Asclepiadaceae). 

All of these host plants (except Brachystelma) possess obvious white latex, which is 
probably ingested by the Euploea larva. Australian Hoya australis and certain popula- 
tions of Sarcostemma australe s.l. have been found to be highly toxic on ingestion by 
domestic livestock, with as yet unsatisfactorily determined chemical components of the 
latex implicated (Everist 1981, Poisonous plants of Australia, Angus & Robertson, Syd- 
ney). Presumably the Euploea larvae are not affected by this toxic principle. 

The chemical composition of latex and leaf surface waxes of different species of Hoya 
differs in the proportions and presence of various triterpenols and their esters (Baas & 
Niemann 1979, Planta Medica 35:348-353; Baas, Warnaar & Niemann 1981, Acta. Bot. 
Neer]. 30:257-263; Warnaar 1984, Phytochemistry 23:1049-1053). As the composition 
of these components appears to be species specific, it would be of interest to investigate 
whether or not selective Euploea feeding on different species of Hoya is correlated with 
these chemical differences in the host plants. 


P. I. ForsTER, Botany Department, University of Queensland, St. Lucia, Queensland 
4067, Australia. 


Received for publication 27 May 1986; accepted 8 August 1986. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 356-357 


BOOK REVIEW 


THE BUTTERFLIES OF NORTH AMERICA: A Natural History and Field Guide, by James 
A. Scott. 1986. Stanford University Press, Stanford. 583 pp., 64 color plates. $49.50. 


To generations of American collectors, Holland’s Butterfly Book was the butterfly 
book. For eastern North America, it was superseded in 1951 by Klots’ Field Guide. No 
equivalent work appeared in the West until 1986, when Tilden & Smith’s Field Guide 
in the Peterson series finally came out. The uneven and flawed Howe Butterflies of 
North America (1975) was supposed to be an update of Holland, but never really “caught 
on.” Meanwhile, there had been a proliferation of regional treatments, from Shapiro’s 
Butterflies of the Delaware Valley (1966) and Pyle’s Watching Washington Butterflies 
(1974) through Emmel & Emmel on southern California, Dornfeld on Oregon, Ferris 
and Brown on the Rockies, and most recently Opler & Krizek on the eastern fauna again. 
Scott’s new book, ambitiously also titled The Butterflies of North America, is another 
attempt to encompass the whole fauna as Holland had done. The result is a visually 
striking book, and one that belongs in every lepidopterist’s library. But it is not without 
its problems. 

While regional treatments were proliferating, taxonomic problems were, too. A great 
deal of ink, and many hurt feelings, have been spent in wrangling over the “correct” 
names of North American butterflies. The controversy is too recent as history to require 
recapitulation here, but in a nutshell, most of the names familiar for two or three gen- 
erations were altered by a combination of generic splitting and, later, a zealous attempt 
to make all specific epithets agree in gender with their current genera. The latter rep- 
resented a strict interpretation of the Code. The former was voluntary, and represented 
an interpretation of the concept of the genus which was in tune with what prominent 
amateurs in Europe were doing but, in the opinion of many, was inadequately justified 
in biology. There are no external standards of right or wrong here; the concept of the 
genus is thoroughly nebulous and must forever be. Meanwhile, collectors need to put 
names on their beasties. Anyone who tries to use Scott and almost any other recent book 
together, and who is not already taxonomically sophisticated, may decide to give up 
butterflies altogether and take up Latin blank verse instead. 

The reason for this is that Scott is diametrically opposed to the trend of the times. He 
is a ‘lumper.’ He lumps at both generic and specific levels; thus one finds most of the 
familiar generic names restored (which I applaud)—but sometimes it becomes rather 
difficult to figure out what species is meant. His judgment is very likely to be correct 
(as, for example, on putting ferrisi as a subspecies of Lycaena rubidus, and ignoring the 
mysterious Johnson Mitouras altogether) in a majority of cases, but the justification for 
it is often unstated. In other cases (putting all of the Cupressaceae-feeding Mitoura in 
North America except hesseli in one huge species, gryneus, for example) the lumping is 
so ambitious that it is very likely wrong—and certain to inspire much indignation. 

As a specimen problem case, consider the Hesperia “comma complex.” I am not 
convinced that the Nearctic taxa should be put as subspecies of the Palearctic comma at 
all (ditto the Coenonympha “tullia complex’), but be that as it may, there is a generally 
accepted set of subspecies in western North America, described, defined, and mapped 
in some detail in MacNeill’s monograph, and not substantially altered in any taxonomic 
work appearing since except for being put under comma, instead of harpalus as a specific 
epithet. We happen to be working very hard on the genetics of the northern California 
members of this complex, so I turned to Scott’s treatment—and was flabbergasted. Instead 
of the ten subspecies in Tilden and Smith, Scott recognizes eight, plus a new unnamed 
one. One of the eight is his own, and does not appear in Tilden and Smith (oroplata). 
The treatment of the California populations is revolutionary: Scott puts ssp. manitoba 
“south in the Cascades to northern California,” while Tilden and Smith, following 
MacNeill, have it south only to northern Washington and Wyoming. Ssp. oregonia dis- 
appears into manitoba without a word. Ssp. yosemite is given by Scott as ranging over 
“west slope of the Sierra Nevada, southern California, and the Inner Coast Range’ — 


VOLUME 40, NUMBER 4 RP ws 


thus silently incorporating both tildeni and leussleri. The figured specimen shows a 
phenotype never remotely approached on the Sierran west slope, whence comes the type. 
That is not surprising; it is from the Bay Area (Santa Clara County)! And so on. 

And one could go on, easily, for many pages. But the message can be defined succintly: 
Scott may be right a lot of the time, but a popular book—a “field guide and natural 
history’ —is not the appropriate place to shake up formal taxonomy like this. There is 
no one to enforce taxonomic decisions except journal editors, and they are not of any 
one mind. What is the collector to do? 

All this said, I must admit that Scott’s book is beautiful. The color plates are all 
photographic, and are by far the most useful ever produced in this country for identifi- 
cation purposes, even if one is completely confused about the names. This is particularly 
true of the skippers, most of which have never been reproduced as color photographs 
before. The specimens in the plates are identified by the number of the species in the 
text. This makes it easy to reference from plate to text. To go the other way is to hunt 
and peck, because the specimens are not arranged in numerical order. 

The host plant information is copious, not referenced, and apparently subject to some 
winnowing (and occasional critical comment), but inadequately qualified as to local 
specialization vs. species-wide breadth. The range maps include much recent informa- 
tion, and they correct a few errors that appeared in the Opler & Krizek book. The text 
is full of biological information, mostly attributed, which is unfamiliar and much of 
which has certainly never been published before. There is a splendid and entirely novel 
section on larval morphology, complete with setal maps and a brand-new key to first- 
instars. There is a “hostplant catalogue’ at the back which enables the reader unarmed 
with the two volumes of Tietz to go from the identity of a plant to possible identities of 
the larva found eating it (but of course, this presupposes that it is a butterfly, not a moth, 
larva). 

In short, an amazing, impressive, infuriating book that will give us all much to argue 
about for decades. I am reminded of the lovers in a Noel Coward play who cannot live 
with or without each other. As I continue to mine this book for goodies and to fume 
over this or that piece of lumping or offhand undocumented zinger, I am thankful that 
I no longer maintain a private collection and that the responsibility for curating our 
institutional one is not mine. 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Journal of the Lepidopterists’ Society 
40(4), 1986, 358-360 


INDEX TO VOLUME 40 


(New names in boldface) 


Aberration, 68 
Ackery, P. R., 77 
Allyn, A. C., obituary, 75 
Allyson, S., 315 
Amateur, 1, 247 
Announcements, 
key words, 126 
manuscript dating, 58 
Arctiidae, 131, 206 
Arnold, R. A., 238 
Artogeia rapae, 79 
Asclepias, 255 
Austin, G. T., 55 
Autographa 
buraetica, 158 
ottolenguti, 158 
Ballmer, G. R., 127 
Battus philenor, 191, 348 
Beck, A. F., 351 
Beirne, B. P., 240 
Bergomaz, R., 131 
Bian, Z., 36 
Biology, 23, 27, 36, 98, 264, 289, 304, 315 
Blum, M. S., 36 
Book reviews, 72, 74, 123, 240, 288, 356 
Boppreé, M., 131 
Bowers, M. D., 214 
British Pyralid Moths, book review, 240 
Brower, L. P., 164, 255 
Brown, F. M., 7 
Brown, R. L., 327 
Butler, L., 289 
Butterflies East of the Great Plains, book 
review, 288 
Butterflies of Europe, vol. 1, book review, 
123 
Callaghan, C. J., 93 
Calvert, W. H., 164 
Carterocephalus silvicolus, 351 
Catastega, 327 
Ceanothus, 350 
Chionodes spp., 298 
Citheronia splendens sinaloensis, 264 
Clarke, J. F. G., 106 
Cleoeromene, 271 
Coutsis, J. G., 97 
Covell, C. V., 128 
Crabtree, L., 206 
Crambinae, 107, 271, 315 
Cryan, J. F., 242 
Danaidae, 20, 67, 164 
Danaus plexippus, 67, 164, 255 
Datana major, 318 


Defense, 191 
DeVries, P. J., 124 
Dichrorampha broui, 322 
Diet, 131 
Diptychophora diasticta, 107 
Dirig, R., 242 
Dix, M. E., 298 
Downey, J. C., 288 
Eacles oslari, 264 
Ectomyelois muriscus, 64 
Emergence, 64, 314, 350 
Epargyreus spanna, 59 
Epidemia mariposa, 127 
Epidromia fergusoni, 8 
Epinotia 

sotipena, 327 

vertumnana, 327 
Epiplemidae, 353 
Eubaphe medea, 353 
Euchloe guaymasensis, 188 
Eucosma rosaocellana, 322 
Eumaeini, 138 
Euploea core corinna, 354 
Euprosterna lacipea, 353 
Euremia nicippe, 130 
Exoteleia anomala, 23 
Fales, H. M., 36 
Fasoranti, J. O., 350 
Ferguson, D. C., 353 
Ferris, C. D., 247 
Forster, P. I., 354 
Garnett, W. J., 351 
Garrison, R. W., 74 
Gaskin, D. E., 107, 271 
Gelechiidae, 23, 298 
Geometridae, 289, 304, 353 
Geometrinae, 304 
Gnophaela latipennis, 206 
Godfrey, G. L., 206 
Greig, N., 124 
Growth, 214 
Hawkeswood, T. J., 347 
Hemileuca 

diana, 27 

grotei, 27 

lucina, 214 

maia, 214 
Hendricks, P., 129 
Henson, P. M., 79 
Hesperiidae, 55, 59, 351 
Higgins, L. G., obituary, 77 
Holland, R., 68 
Hosts, 127, 354 


VOLUME 40, NUMBER 4 


Icaricia icarioides lycea, 68 


Immature stages, 20, 27, 264, 289, 304, 315, 


318 


Incaeromene subuncusella, 271 


Jacobson, M., 298 
Johnson, K., 59, 65 


Key words, announcement, 126 


Knudson, E. C., 322, 353 
Lafontaine, J. D., 158 
Limacodidae, 69, 353 
Lycaenidae, 68, 127, 138, 238 
Lycorea pieteri, 20 

Malcolm, S. B., 255 


Manuscript dating, announcement, 58 


Mating, 238 

Matusik, D., 59, 65 
Medley, M. E., 128 
Microcrambus elegans, 315 


Milkweed Butterflies, book review, 72 


Miller, G. L., 318 
Miller, J. Y., 75 
Millers L. D., 1, 72, 75 
Miller, W. E., 218 


Molybdogompha polymygmata, 3538 
Monograph of the Birdwing Butterflies, 


book review, 74 
Munree, E., 241 
Mynes geoffroyi guerini, 347 
Natural history, 206 


Nemoria glaucomarginaria, 304 


Nemoriini, 304 
Neoeromene parvipuncia, 271 


Neophasia menapia menapia, 314 


Noctuidae, 8, 124, 128, 158 


Nordeuropas Pyralider, book review, 241 


Notodontidae, 318 
Nymphalidae, 347, 354 
Obituaries, 75, 77, 242 
Ocalaria, 124 
Olethreutinae, 218, 322 
Opler, P. A., 188 
Overwintering, 164 
Oviposition, 130, 255, 348 
Pammene medioalbana, 322 
Papaj, D. R., 348 
Papilio 

anchisiades, 36 

xanthopleura, 65 
Papilionidae, 36, 65, 191, 348 
Parasitization, 347 
Pareromene, 271 
Pericopinae, 206 
Phigalia strigateria, 289 
Phobetron hipparchia, 69 
Phoebis 

editha, 97 

sennae, 97 


Pieridae, 79, 97, 130, 188, 314 
Pieris rapae, 79 
Plagens, M. J., 130 
Plusiinae, 158 
Polychrysia morigera, 128 
Porter, A. H., 304 
PratiiG F027 
Predation, 129, 191, 351 
Presidential addresses, 

1984, 1 

1986, 247 
Psamathia placidaria, 353 
Pseudexentera 

hodsoni, 218 

knudsoni, 218 

oreios, 218 

sepia, 218 

spp., 218 

vaccinii, 218 
Pyralidae, 64, 107, 271, 315 
Pyrgus 

albescens, 55 

communis, 55 
Reakirt, T., article about, 7 
Rearing, 131 
Restinga, 93 
Retinia 

metallica, 298 

spp., 298 
Rhyacionia spp., 298 
Riodinidae, 93 
Riodininae, 238 
Robbins, R. K., 79, 106, 138 
Roosting, 124 
Rozycki, R., 65 
Rupert, L. R., obituary, 242 
Saturnia walterorum, 54 
Saturniidae, 27, 54, 214, 264 
Shapiro, A. M., 356 
Shields, O., 123 
Small-Nicolay collection, 106 
Smiths Vv, 67 
Solis, M. A., 8 
Stamp, N. E., 191, 214 
Steneromene, 107 
Steruliaceae, 64 
Stevens, R. E., 23 
Synargis brennus, 98 


359 


The Butterflies of North America, book 


review, 356 
Theobroma simiarum, 64 
Tischeria immaculata, 350 
Tischeriidae, 350 
Tortricidae, 218, 298, 322, 327 
Trapping, 298 
Trichonis, 188 
Tuskes, P. M., 27, 264 


360 


Ultrastructure, 318 
Valverde, M. D., 54 
Wetherbee, D. K., 20 
Williams, M. L., 318 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Wolfe, K. L., 54 
Young, A. M., 36, 64, 69 
Young, R. M., 314 


Date of Issue (Vol. 40, No. 4): 11 March 1987 








EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
BoYcE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
THEODORE D. SARGENT, ROBERT K. ROBBINS, ROBERT C. LEDERHOUSE 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Short manuscripts concerning new state records, current events, and notices 
should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence, KS 66044 
U.S.A. Journal contributors should prepare manuscripts according to the following in- 
structions. 

Abstract: An informative abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article's content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name with author. Mea- 
surements should be given in metric units; times should be given in terms of the 24- 
hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. References 
to footnotes should be numbered consecutively, and the footnotes typed on a separate 
sheet. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 196la, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In general notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-—216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30). 

Illustrations: Only half of symmetrical objects such as adults with wings spread 
should be illustrated, unless whole illustration is crucial. Photographs and drawings should 
be mounted on stiff, white backing, arranged in the desired format, allowing (with 
particular regard to lettering) for reduction to their final width (usually 4% inches). 
Illustrations larger than 8% x 11 inches are not acceptable and should be reduced pho- 
tographically to that size or smaller. The author’s name, figure numbers as cited in the 
text, and an indication of the article’s title should be printed on the back of each mounted 
plate. Figures, both line drawings and halftones (photographs), should be numbered 
consecutively in Arabic numerals. The term “plate” should not be employed. Figure 
legends must be typewritten, double-spaced, on a separate sheet (not attached to the 
illustrations), headed EXPLANATION OF FIGURES, with a separate paragraph devoted to 
each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the 
proofs. A page charge of $60 per full Journal page is applicable to all authors. Unfunded 
authors may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 





CONTENTS 


PRESIDENTIAL ADDRESS, 1986: UNEXPLORED HORIZONS—THE ROLE 
OF THE AMATEUR LEPIDOPTERIST. Clifford D. Ferris ......... 
SELECTIVE OVIPOSITION BY MONARCH BUTTERFLIES (Danaus 
PLEXIPPUS L.) IN A MIXED STAND OF ASCLEPIAS CURASSAVICA 


247 


L. AND A. INCARNATA L. IN SOUTH FLORIDA. Stephen B. — 


Malcolm t+ Lincoln P. Brower ca i 


BIOLOGY AND IMMATURE STAGES OF CITHERONIA SPLENDENS SI- 


NALOENSIS AND EACLES OSLARI IN ARIZONA (SATURNIIDAE). 


Paul M. Tuskes oo 


NEW GENERA FOR THE NEOTROPICAL “PAREROMENE’ speemea 
(PYRALIDAE: CRAMBINAE). David E. Gaskin coe 


BIOLOGY AND DESCRIPTION OF IMMATURE STAGES OF PHIGALIA 
STRIGATERIA (MINOT) (GEOMETRIDAE). Linda Butler _. 


TRAP PREFERENCES OF RETINIA METALLICA AND SEASONAL FLIGHT 
BEHAVIOR OF RETINIA SPP., RHYACIONIA SPP. (TORTRICIDAE), 
AND CHIONODES SPP. (GELECHIIDAE) IN THE DAKOTAS. 
Mary Ellen Dix & Martin Jacobson ee 


LIFE HISTORY OF NEMORIA GLAUCOMARGINARIA (BARNES & 
McCDUNNOUGH) AND LARVAL TAXONOMY OF THE TRIBE NE- 
MORIINI (GEOMETRIDAE: GEOMETRINAE). Adam H. Porter 

SOD WEBWORMS: THE LARVA OF MICROCRAMBUS ELEGANS (CLEM.) 
(PYRALIDAE: CRAMBINAE). Suzanne Allyson ic iccccccceueennn 

ULTRASTRUCTURE OF THE EGG OF THE AZALEA CATERPILLAR, 
DATANA MAJOR GROTE & ROBINSON (NOTODONTIDAE). 
Gary L. Miller G Michael L. Williams 0) oe 

NEW SPECIES OF OLETHREUTINE MOTHS (TORTRICIDAE) FROM 
TEXAS AND LOUISIANA. Edward C. Knudson 7 ae 


RESURRECTION OF CATASTEGA CLEMENS AND REVISION OF THE 
EPINOTIA VERTUMNANA (ZELLER) SPECIES-GROUP (TORTRI- — 


CIDAE: OLETHREUTINAE). Richard L. Brow? cece 


GENERAL NOTES 


Mass emergences of the pine white, Neophasia menapia menapia (Felder & 
Felder), in Colorado (Pieridae). Romald M. YOUt gy -neesecc.ecccecssseccccseesseeesesseneneeeee 
Mynes geoffroyi guerini Wallace (Nymphalidae) parasitized by a tachinid fly. 
T. J. Hawkeswood (xij 


An oviposition “mistake” by Battus philenor L. (Papilionidae). Daniel R. 
PQ Pay) ices tN ye EN EST TO 


Emergence of Tischeria immaculata (Braun) (Tischeriidae) from leaves of 
Ceanothus griseus L. J. Olaniyan Fasoranti 


Unusual predator damage to Carterocephalus silvicolus (Meigen) (Hesperi- 
idae). . Andrew F. Beck ¢ William J. Garnett 


Four new United States records of moths from Texas. Douglas C. Ferguson 
d> Edward C. Knudson 20 


New Host records for Euploea core corinna (Macleay) (Nymphalidae). P. I. 
Forster 


Book REVIEWS 
Butterflies East of the Great Plains. 
The Butterflies of North America: A Natural History and Field Guide............ 


INDEX TO VOLUME 40 





(255 


264 
271 


289 


298 


304 


315 


318 


322 


2 Syn a EES eS RE es zB. 


Volume 41 1987 . Number 1 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





16 April 1987 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


DoucGLas C. FERGUSON, President GERARDO LaAMas M., Vice President 
Jerry A. POWELL, President-Elect EBBE SCHMIDT NIELSEN, Vice 
CLIFFORD D. FERRIS, Immediate Past President President . 

OLAF H. H. MIELKE, Vice President JAMES P. TUTTLE, Treasurer 


RICHARD A. ARNOLD, Secretary 


Members at large: 


BoyYcE A. DRUMMOND III MIRNA M. CASAGRANDE M. DEANE BOWERS 
JOHN LANE EDWARD C. KNUDSON RICHARD L. BROWN 
ROBERT K. ROBBINS FREDERICK W. STEHR PAUL A. OPLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 


Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopierists’ Society, to: James P. Tuttle, Treasurer, 
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian 
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007- 
4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Secretary, 
50 Cleaveland Rd., #3, Pleasant Hill, CA 94523-3765, U.S.A. 


To obtain: 

Back issues of the Journal and NEWS (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society’’) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, CA 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los 
Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, CA and additional 
mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % 
Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 


Cover illustration: Semilooping larva of the strange noctuid Phyprosopus callitrichoides 
on Smilax. Sketch by Mark Klingler, Carnegie Museum of Natural History. Suggested 
by John E. Rawlins. 


JouRNAL OF 
THe LeEPIDOPTERISTS’ SOCIETY 


Volume 41 1987 Number 1 


Journal of the Lepidopterists’ Society 
41(1), 1987, 1-12 


STATUS AND HABITATS OF POTENTIALLY ENDANGERED 
LEPIDOPTERA IN OHIO 


JOHN A. SHUEY, ERIC H. METZLER, DAVID C. IFTNER, 
JOHN V. CALHOUN, JOHN W. PEACOCK, REED A. WATKINS, 
JEFFREY D. HOOPER AND WILLIAM F. BABCOCK 


The Ohio Lepidopterists, 1241 Kildale Square North, 
Columbus, Ohio 43229 


ABSTRACT. The status of eight species that are potential candidates for addition to 
the U.S. list of endangered species was assessed in 1985. Two of these, Phyciodes batesii 
(Nymphalidae) and Acronicta albarufa (Noctuidae), are known only from literature 
records, and their occurrence in Ohio is unverified. Three species, Neonympha mitchellii 
(Satyridae), Catocala marmorata (Noctuidae), and Catocala pretiosa (Noctuidae), have 
not been collected in Ohio for more than 30 years. Three species are extant in Ohio: 
Lycaeides melissa samuelis (Lycaenidae), Speyeria idalia (Nymphalidae), and Erythoecia 
hebardi (Noctuidae). Further investigations into the ecological requirements of all the 
species are suggested, as well as habitat manipulations and acquisitions to insure their 
continued survival in Ohio. Special emphasis should be placed upon melissa and hebardi 
because both are limited to small geographic areas in the State. Speyeria idalia seems 
secure in unglaciated Ohio but has undergone a decline in glaciated areas. Conservation 
efforts for this species should be concentrated on the isolated populations in glaciated 
Ohio. 


Additional key words: surveys, conservation. 


Eight species of Ohio Lepidoptera, four butterflies and four moths, 
have been identified as being potentially threatened or endangered 
(Anonymous 1984). The species are: 1) Lycaeides melissa samuelis 
Nabokov (Lycaenidae), 2) Speyeria idalia (Drury) (Nymphalidae), 3) 
Phyciodes batesii (Reakirt) (Nymphalidae), 4) Neonympha mitchellii 
French (Satyridae), 5) Acronicta albarufa (Grote) (Noctuidae), 6) Ca- 
tocala marmorata W. H. Edwards (Noctuidae), 7) Catocala pretiosa 
Lintner (Noctuidae), and 8) Erythoecia hebardi Skinner (Noctuidae). 

Concerning the butterflies, P. batesii is known only from literature 
records, and its occurrence in Ohio is unverified. Neonympha mitchellii 
has not been collected in Ohio for more than 30 years. These species 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


may occur in Ohio, but unless they are rediscovered, no action can be 
directed toward their preservation. Lycaeides melissa samuelis and S. 
idalia are extant in Ohio, and further investigation into their ecological 
requirements is necessary to insure their continued survival in the State. 

Of the four species of moths, only one, E. hebardi, has recently been 
collected in Ohio. One, A. albarufa, is known only from literature 
records, and the other two, C. marmorata and C. pretiosa, have not 
been seen in Ohio since the end of the nineteenth century. Based on 
the written reports and the numbers of extant specimens, it can be 
deduced that C. pretiosa was not rare, whereas C. marmorata was. 

The brevity of some of the species reports to follow here is evidence 
of how little is known about them. This is perhaps due to their rarity, 
but dearth of written information is typical of economically unimpor- 
tant Lepidoptera. Although many collectors have “local” knowledge, 
it is rarely written down. Specimens in collections provide some clues, 
but researchers frequently work without much information. 

Information on species potentially threatened and endangered in 
Ohio has not been adequately compiled. Many areas in Ohio provide 
habitats for potentially threatened or endangered species of Lepidop- 
tera. Data on these species and their habitats are needed to enhance 
our ability to make biologically sound policy and management decisions 
concerning the species and their habitats. 

The Ohio Lepidopterists, an organization dedicated to advancing the 
scientific knowledge cf Lepidoptera, conducted a one-year study of the 
habitats and plant associations of the target Lepidoptera species, their 
presence or absence in selected habitats, and their historical occurrence 
in Ohio. The following is a summary of findings. 


SPECIES ACCOUNTS 


Lycaeides melissa samuelis 


Historical distribution. The eastern subspecies samuelis occurs in scattered colonies in 
the Great Lakes area and the Northeast (Opler & Krizek 1984). This insect, the Karner 
blue, has long been known from Ohio (Rawson & Thomas 1939, Nabokov 1949, Forbes 
1960, Price 1970, Opler & Krizek 1984). In recent years, it has been recorded only from 
an area adjacent to the Schwamberger Preserve in Lucas Co. (Fig. 1). A single record 
from Summit Co. (Albrecht 1982) was based on a misidentification. 

Habitat and plant associations. Lycaeides melissa samuelis inhabits sandy pine barrens, 
oak openings, lakeshore dunes and sandy pine prairies. These habitats must support the 
lupine, Lupinus perennis L., the only known larval hostplant of samuelis. Lupinus 
perennis, itself considered “potentially threatened” in Ohio (Cooperrider 1982), requires 
periodic fire or other disturbances to compete with woody plants (Dirig & Cryan 1976, 
Miller 1979). Lupine grows in sandy soils, and is important in stabilizing open sand. 
However, as the soil is stabilized, trees become established, and herein lies the problem 
for the continued survival of the Karner blue. Apparently, the butterfly will not utilize 
shaded lupine plants. Originally, natural wildfire was probably an important factor in 


VOLUME 41, NUMBER 1 3 















Bees) 
ros 


i 






A L. melissa samuelis 


@ N. mitchellii 


Fic. 1. Distribution of Lycaeides melissa samuelis and Neonympha mitchellii in 
Ohio, based on examined specimens. 


controlling trees that encroached the prairies and other openings. The recent advent of 
fire prevention has resulted in many of the prairies of the area becoming overgrown with 
shrubs. Some of these areas have become quickly forested. One of the many results of 
this has been the near extinction of the Karner blue in Ohio. The last known population 
could easily become extinct despite efforts to protect it. 

Current distribution in Ohio. Presently, a small area in and near the Schwamberger 
Preserve harbors the only known natural population of the Karner blue remaining in 
Ohio. Several other potential habitats in Lucas Co. have been surveyed by the Ohio 
Lepidopterists and others in recent years. Only one other population of samuelis was 
discovered, an introduced small colony in Oak Openings Metropark near Toledo. Scattered 
populations of lupine were discovered at a few sites in Lucas Co. (Campbell State Nature 
Preserve, and Oak Openings Metropark system) and in Henry Co., but no samuelis were 
seen at any of these locations. 


Discussion. Presently, the outlook for the Karner blue in Ohio is not 
favorable. The butterfly has been virtually eliminated from the Oak 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





{ @ post-1970 
~ © pre-1970 


7x literature 





Fic. 2. Distribution of Speyeria idalia in Ohio, based on examined specimens and 
literature records. Line indicates S limit of Wisconsin glacier. 


Openings area, very likely as a result of the forestation of the original 
sandy prairies. Lands in and near the Schwamberger Preserve contain 
a small population of samuelis, but these areas will require intense 
management if the population of samuelis (and other rare butterflies 
and plants) is to survive. 

The Karner blue (and two other rare Ohio butterflies, Incisalia irus 
(Godart) and Erynnis persius Scudder) should benefit from habitat 
manipulations that would increase the local distribution and density of 
Lupinus at the Preserve. Large stands of Lupinus currently exist in at 
least two sites within the Preserve. One site seems ideal for manipu- 
lations to establish the plant in the nearby sandy ex-agricultural area. 
It is likely that prescribed burns could be utilized to enhance the prairie 


VOLUME 41, NUMBER 1 5 


aspects of the Preserve. Samuelis may also benefit from the thinning 
of shrubby vegetation within the Lupinus stands. Recent bulldozing of 
aspen from land in the Preserve may have inadvertently benefited 
samuelis by removing shade from existing lupine. Samuelis was not 
observed in this area in July 1983, but was frequent in May and July 
1984. 


Speyeria idalia 


Historical distribution. Speyeria idalia, the regal fritillary, ranges from N New England 
W to S North Dakota, S across the N half of the U.S. to E Colorado and Montana (Opler 
& Krizek 1984). This species was first recorded in Ohio in 1854 (Kirtland 1854). Histor- 
ically, idalia has been most often recorded from the E half of Ohio, although early records 
indicate it was once common in NW Ohio (Dury 1878, Bubna 1897, Hine 1898a, 1898b, 
Bales 1909, Henninger 1910, Wyss 1930, 1932). Albrecht (1982) reported that Ohio records 
include much of the State. It is ironic that there are so few records for idalia in the 
- W-central counties of Ohio. This region was once a vast mesic prairie, which should have 
supported large populations of idalia. Perhaps the paucity of records from W-central 
counties reflects the rapid degradation of idalia habitat following conversion of prairies 
to agricultural land in the mid-1800’s, which occurred prior to active collecting in Ohio. 
In recent years, most collections have been in the SE quarter of the State (Fig. 2). 

Habitat and plant associations. The regal fritillary is a butterfly of tall-grass prairie in 
the Midwest, but is found in other open grassy situations elsewhere. In the East, it is 
found in damp meadows or pastures with boggy or marshy areas, but it inhabits dry 
mountain pastures in some areas. The reported primary larval host of idalia is bird’s foot 
violet (Viola pedata L.); other Viola species may be utilized. Pedata is extremely rare in 
SE Ohio (Cusick & Silberhorn 1977), so another violet may be the larval host there. Adults 
commonly nectar on thistles and milkweeds, along with red clover in pastures. 

Curreni distribution in Ohio. The range of idalia in Ohio has apparently diminished 
in recent years. Most recent records are from SE counties, although there are some records 
for a few N-central counties since 1970. It has not been recorded in recent years from 
the NW part of the State. This decline in numbers is not limited to Ohio, as this species 
appears to have declined precipitously in many areas, and is common only in the few 
remaining untilled areas in the prairie States (Hammond & McCorkle 1983). In most 
states, idalia is now present only in fragmented populations, and has been extirpated 
from large regions where it was once common, such as parts of the Ohio Valley and the 
N Midwest (Opler & Krizek 1984). Its disappearance is likely due to destruction of native 
prairie habitat, and along with it, destruction of larval host violets. 


Discussion. Speyeria idalia could probably be preserved in glaciated 
Ohio by promoting a return of suitable areas to original wet prairie 
habitats through land purchases or specific land management practices. 
Presently, this species appears to be unthreatened in unglaciated Ohio. 
However, it is conceivable that the general decline of idalia indicates 
the future for this species in SE Ohio, and thus its status should be 
periodically monitored. An effort should be made to determine the 
larval host plant in SE Ohio. 


Phyciodes batesii 


Historical distribution. The tawny crescent, Phyciodes batesii, ranges from S Quebec 
and Ontario S to Pennsylvania and Michigan, and W to Nebraska and Colorado. It has 


6 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


been found in isolated colonies farther S in the Appalachians in Virginia, Kentucky, North 
Carolina, and Georgia (Opler & Krizek 1984). 

All the numerous literature records for this species in Ohio (Bales 1909, Henninger 
1910, Wyss 1932, Studebaker & Studebaker 1967) are considered dubious; existing spec- 
imens thought to be batesii have been determined to be Phyciodes tharos (Drury), a 
common Ohio species. 

Habitat and plant associations. In the N part of its range, batesii is found in low-lying 
moist meadows or pastures. In its S distribution, it is found on the tops of dry, rocky 
bluffs above rivers, or on dry hillsides or rocky upland pastures, usually in association 
with Andropogon grass. The larval foodplant is wavy-leaved aster (Aster undulatus L.) 
and possibly other true asters (Opler & Krizek 1984). 

Current distribution in Ohio. There is no evidence that this species currently exists in 
Ohio. 


Diseussion. Cusick and Silberhorn (1977) record A. undulatus, the 
known larval foodplant, in 21 of 83 SE Ohio counties. These areas and 
others where A. undulatus is native should be thoroughly explored for 
batesii. 


Neonympha mitchellii 


Historical distribution. Mitchell’s satyr, Neonympha mitchellii, displays a disjunct 
distribution. The only known localities are in New Jersey (Rutkowski 1966), South Car- 
olina, Indiana, Michigan, and Ohio. Midwestern sites occur in a limited area characterized 
by glacial till topography and calcareous springs. 

In Ohio, mitchellii was first recorded in Streetsboro Fen in Portage Co. (Pallister 1927) 
(Fig. 1). Holland (1931) and Macy and Shepard (1941) also cited mitchellii in Ohio. 
Pallister found mitchellii abundant in Streetsboro Fen on 4 July 1925 and 10 July 1926. 
He described the area as a several-hundred-acre peat swamp, but found that mitchellii 
was restricted to approximately one acre of “sedge meadow’ surrounded by tamarack 
and maple. According to McAlpine et al. (1960), mitchellii was last reported from the 
area on 19 June 1950. By 1954, most of Streetsboro Fen had been converted to a truck 
farm. 

Habitat and plant associations. Although the literature reports that mitchellii occurs 
in bogs, all of the habitats described are clearly fens (Shuey 1985) (bog fens in the 
terminology of Stuckey & Denny 1981). Fens occur over alkaline springs on deposits of 
peat, and are dominated by sedges (bogs are acidic and are dominated by mosses in the 
genus Sphagnum) (Pringle 1980). Reliable indicators of potential mitchellii habitats 
include tamarack, poison sumac, shrubby cinquefoil and abundant sedges. Mitchellii has 
a strong preference for flying in open stands of tamarack, especially along stream banks. 

Mitchellii has been reared on several Carex species (McAlpine et al. 1960), but it is 
not known what species are utilized in the natural habitat. Mitchellii is usually closely 
associated with narrow-leaved Carex, probably C. stricta Lam. or C. aquatilus Wahlenb. 

Current distribution in Ohio. Today, there remains little habitat in Ohio resembling 
the habitats that support viable colonies of mitchellii in Indiana and Michigan. Gott Fen 
State Nature Preserve, located within Streetsboro Fen, is primarily a shrubby cinquefoil 
meadow, but some limited areas support lush Carex openings with which mitchellii is 
associated. Mitchellii generally flies in lush sedge meadows adjacent to such areas, but at 
Streetsboro these are the areas that seem most heavily disturbed. Most sedge meadows in 
Streetsboro Fen support the broad-leaved sedge, Carex lacustris Willd., or rushes, Scirpus 
spp., not the narrow-leaved species with which mitchellii is typically associated. 

On 29, 30 June, and 6, 7 July 1985, several members of The Ohio Lepidopterists 
surveyed for mitchellii at four fens in Portage Co. (Wingfoot Lake, Mantua Swamp, 
Frame Lake Bog/Herrick Preserve and Gott Fen State Nature Preserve), one fen in Stark 
Co. (Timken Bog), and two fens in Summit Co. (Standard Slag Bog and Nimisila Bog 


VOLUME 41, NUMBER 1 7; 


Meadow). Mitchellii was not located in these surveys and, because mitchellii is usually 
common when found, and the flight period was well covered in these surveys, it was 
concluded that mitchellii probably is not present at these sites. 


Discussion. If mitchellii is to be rediscovered in Ohio, efforts should 
be concentrated in habitats similar to the original Streetsboro Fen. The 
fen at Herrick Nature Preserve, just S of Streetsboro Fen, lacks suitable 
sedge meadows, but as far as known, no one has searched Tinkers Creek 
between these two areas. Many other fens (Mantua Swamp, Kick Fen, 
others) in NE Ohio fit the general habitat description. Unfortunately, 
fire suppression has allowed many of these areas to become overgrown 
with species of dogwood, reducing the amount of suitable sedge mead- 
ow. Mantua Swamp and Standard Slag Bog were not completely sur- 
veyed in 1985 because of their large size. If sedge meadows occur in 

-the center of these sites, they deserve a closer look. If ““new’’ fens are 
discovered in the Portage Co. area that contain extensive stands of 
tamarack, they, too, should be sampled for mitchellii. 

In NW Ohio, the only likely habitat that remains in undisturbed 
condition is Mud Lake in Williams Co. Mitchellii does not occur at 
Mud Lake, and probably has not occurred there in recent times (Price 
1970). However, Mud Lake is very similar botanically to nearby Cedar 
Lake fen in NE Indiana, which does support a colony of mitchellii. 


Catocala marmorata 


Historical distribution. The marbled underwing, Catocala marmorata, is one of the 
rarer taxa in the genus. It has not been seen in Ohio since late in the nineteenth century 
(Dury 1876, 1878, Pilate 1882), and has always been rare over its entire range (Holland 
1903, Barnes & McDunnough 1918, Forbes 1954, Sargent 1976). Sargent (pers. comm.) 
indicated that marmorata was collected more regularly in recent years. Collectors in 
Kentucky have taken about a dozen specimens in the past 10 years; compared to previous 
years, this would seem a population explosion. 

All records for Ohio are from the S half in Hamilton, Montgomery, and Franklin 
counties (Fig. 3). These records were made by very active collectors (Charles Dury, 
Cincinnati; George Pilate, Dayton; and W. N. Tallant, Columbus) in the late 1800's. A 
specimen in the Strecker collection in the Field Museum of Natural History, Chicago, is 
simply labeled “S. Ohio, 8-26-[18]76”. Specimens in the Cincinnati Museum of Natural 
History and the Field Museum substantiate the historical occurrence of marmorata in 
Ohio. Because this species is not easily confused with others, literature records can be 
accepted with little hesitation. 

Habitat and plant associations. Habitats and plant associations of marmorata are 
unknown. This species is placed with those of the genus whose caterpillars feed on willow 
and aspen. This placement is supported by overall moth appearance and genitalia (Gall 
1984). 

Current distribution in Ohio. It is difficult to assess the present status of this species 
in Ohio. Although it has not been seen in Ohio for approximately 90 years, there have 
been few active moth collectors in Ohio in that time, and none of them have collected 
in the SW part of the State where marmorata might be found. The species has been 
collected in an upland oak woods in S Kentucky (Loran Gibson, pers. comm.), but 
collecting in SW Ohio in similar habitats has not yielded it. Marmorata certainly is vagile, 
and may well have been a temporary resident or even a vagrant in Ohio, as is probably 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 









ie 
@ 





\ © C. marmorata 


Y @ Cc. pretiosa 
A A. albarufa 
&® E. hebardi 


Fic. 3. Distribution of Catocala marmorata, C. pretiosa, Acronicta albarufa, and 
Erythoecia hebardi in Ohio, based on examined specimens and literature records. 


the case for Connecticut, New York, and New Jersey. Given its rarity, marmorata might 
still be found in Ohio. 


Discussion. A stronger effort to collect in SW Ohio in willow, cot- 
tonwood, and oak habitats is needed. If the supposition is correct that 
the foodplant is salicacious, wet areas of S Ohio, particularly the SW 
portion, will be critical to the survival of this species. If, as indicated 
by some recent captures, the species is associated with oak forests, the 
concern with wet areas will be lessened. With current environmental 
emphases on wetlands as habitats, marmorata may be protected un- 
wittingly. Land changes affecting other forested areas are more difficult 
to control. 


VOLUME 41, NUMBER 1 9 


Catocala pretiosa 


Historical distribution. Catocala pretiosa has been collected in Ohio several times in 
Montgomery and Franklin counties (Fig. 3). All collections were made by W. N. Tallant 
and George Pilate. Pilate and Tallant stopped collecting in Ohio before 1900, when Pilate 
moved to Georgia, and Tallant moved to Richmond, Indiana; pretiosa has not been taken 
in Ohio since. Extant specimens from Ohio are in the Dayton Museum of Natural History, 
Dayton, Ohio, the U.S. National Museum of Natural History, Washington, D.C., and the 
Allyn Museum of Entomology, Sarasota, Florida. Although John and Edward Thomas 
collected extensively in central Ohio (as well as other areas of the State) in the 1930’s, 
they did not collect pretiosa. This is consistent with the distribution reported by Sargent 
(1976), who states, “This moth was not taken by collectors for many years” (from about 
1920 to 1968), and “was often presumed to be extinct’. Schweitzer (1982) discussed recent 
captures of pretiosa, and indicated this species may be restricted to southern New Jersey. 
The New Jersey populations apparently did not exist when pretiosa was being collected 
in other areas, and now that pretiosa is being collected in New Jersey, it is unknown in 
the former locations. 

Habitat and plant associations. Schweitzer (1982) reported rearing pretiosa on Prunus 
maritima Marsh in New Jersey; also (pers. comm.) that he found one larva on Pyrus 
arbutifolia (L.), red chokeberry, in New Jersey in 1986. Other species in this group of 
Catocala also use Pyrus spp. (crabapples), Prunus spp. (plums, cherries) and Crataegus 
spp. (hawthorns) as larval host plants. Prunus maritima and Pyrus arbutifolia do not 
occur in Ohio; therefore, the larval host in Ohio must be some related plant. If the Ohio 
foodplant is a species of Pyrus, Prunus, or Crataegus, the list of possible host species is 
extensive. 

Curreni distribution in Ohio. Three collecting trips were made in central Ohio (SW 
Franklin, Madison counties) in late June-early July 1985 to areas with a concentration 
of possible foodplants. These areas were selected on the basis of the historical distribution 
of pretiosa. General collecting was very good; however, pretiosa was not seen. 


Discussion. Based on current records and Schweitzer’s assumptions, 
we should not expect to find pretiosa in Ohio. 


Acronicta albarufa 


Historical distribution. Acronicta albarufa is recorded from Ohio in only two literature 
records (Bales 1909, Henninger 1910) (Fig. 3). This species is very similar to Acronicta 
ovata Grote, a common species in Ohio. Without verified specimens from Ohio, it is easy 
to imagine that the specimens identified as albarufa may have been ovata. It is not possible 
to exclude albarufa from Ohio’s fauna, however, because the reported species range 
crosses the U.S. from the E coast to New Mexico and Colorado, N to Canada, and S to 
Georgia (Forbes 1954). 

Habitat and plant associations. The apparent habitat requirements of albarufa limit 
the areas where it may be found in Ohio. According to J. G. Franclemont (pers. comm. ), 
the E distribution of albarufa is restricted to sandy soils and habitats consistent with “pine 
barrens’ or “sand barrens’. Once widespread in the E, according to Franclemont, albarufa 
now seems more confined to coastal areas from New Jersey to Cape Cod, and to pine 
barrens near Albany, New York. Schweitzer (pers. comm.) has taken albarufa at two 
locations (Ontario and Massachusetts), and states that the moth occurs in the “‘oak-pine 
forest on the coastal plain”. The oaks are “primarily Quercus velutina, coccinea, stellata 
and alba with some OQ. ilicifolia in the understory”. In Massachusetts, where he has 
collected albarufa, Schweitzer reported “The only tree is Pinus rigida”’ with an understory 
of scrubby oaks that “are well in excess of 99% Q. ilicifolia”. In Grand Bend, Ontario, 
Schweitzer collected albarufa in a “sandy oak (mostly Q. velutina)-pine forest”. 

Current distribution in Ohio. The oak forests of Ohio have been heavily collected in 
the past and albarufa has not been taken. Three collecting trips were carried out in 1985 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in the Oak Openings area of Lucas Co., and the Erie sand barrens in Erie Co. during 
the known flight period which is May through August. General collecting was good, but 
no specimens of albarufa were seen. 


Discussion. Further collecting is needed in suitable habitats, such as 
the Oak Openings area in Lucas Co., and dry ridge tops of S Ohio. For 
now, inclusion of albarufa as part of Ohio’s fauna is tentative. 


Erythroecia hebardi 


Historical distribution. A population of Erythroecia hebardi, one of the rarest moths 
in North America, occurs in Scioto Co., Ohio (Fig. 3). The size of the Ohio population 
is unknown. Originally described in 1917 from two specimens from Hot Springs, Virginia, 
hebardi has not been seen from Virginia since. Until 1984, all known specimens, except 
the types, have come from N-central New Jersey, near Lake Hopatcong and Johnsonburg. 
Before 1984, fewer than 10 specimens were known in collections. Of these, only two, 
those from Johnsonburg, are recent records; all others predate 1930. Nothing is known 
of the life history, and only guesses can be made about its habitat requirements. 

This species was first collected in Ohio at UV light on 26 August 1984 in Shawnee State 
Forest, Scioto Co. The site was a clearing that resulted from a recent clearcutting. More 
collecting in 1985 and 1986 yielded additional specimens, several of which came from 
two additional locations, 1.2 km and 4.2 km S of the first site. The second and third sites 
are also recent clearcuts. 

Habitat and plant associations. In all occurrences of this moth in Ohio, the habitat is 
a recent clearcut area in mature second growth forests in the rugged unglaciated Allegheny 
Plateau of the S part of the State. This is an area of steep hills capped with sandstone 
ridges, and acidic, dry soils. Common forest trees are upland mixed oaks, hickories, maples, 
native pines (Virginia, yellow, and pitch), and yellow poplar. Usual forest groundcover 
plants are several kinds of blueberries, huckleberries, and other members of the heath 
family. Because of the acidic soils, diversity of wildflowers is low (King 1979). 

Adventive species of plants such as clovers, sunflowers, goldenrods, ragweeds, etc., 
quickly invade the clearcut areas to provide considerable diversity of plants not found 
in the forest. All specimens of E. hebardi collected so far have come from clearcut areas 
immediately adjacent to the forest. Larval foodplant and habitat requirements are un- 
known. 

Current distribution in Ohio. To date, this species has been collected only in Scioto 
Co., in S Ohio. 


Discussion. Based on information gathered to date, the continued 
survival of hebardi in Ohio is encouraging. More complete information 
pertinent to its habitat and food requirements is essential. The occur- 
rence of hebardi in Ohio provides a unique opportunity for research 
to discover reasons for its previous rarity. 


CONCLUSIONS 


Most of the species discussed indicate unique ecological situations in 
Ohio. Further investigation into the ecological requirements of all the 
species and their habitats is warranted. The presence of populations 
can and should be used in decisions concerning the preservation of 
unique and endangered habitats. Special emphasis should be placed on 
Lycaeides melissa samuelis and Erythoecia hebardi because both are 
limited to very small geographic areas in Ohio; survey work should be 


VOLUME 41, NUMBER 1 Ll 


directed toward locating additional populations of these species. Spey- 
eria idalia seems secure in unglaciated Ohio, but further research is 
needed to determine larval host plants and the reasons for population 
fluctuations. On the other hand, this species has declined in glaciated 
areas. Conservation efforts for this species should therefore be concen- 
trated on the isolated populations in glaciated Ohio. There should be 
a continuing effort to locate Phyciodes batesii, Neonympha mitchellii, 
Catocala marmorata, Catocala pretiosa and Acronicta albarufa in Ohio. 


ACKNOWLEDGMENTS 


Thanks are due the Ohio Department of Natural Resources, Division of Natural Areas 
and Preserves, for a grant to The Ohio Lepidopterists for this research, and for providing 
access to properties under their jurisdiction. Thanks are also extended to the divisions of 
Parks and Recreation, Forestry, and Wildlife, the Toledo Metroparks, Columbus Metro- 
parks, The Nature Conservancy, and the Ohio Historical Society for information and 
access to properties under their control. We are grateful to J. G. Franclemont, J. D. 
Lafontaine, and D. F. Hardwick for their continuing contributions to research on E. 
hebardi. We thank The Ohio Lepidopterists, curators, collectors, and others too numerous 
to mention who allowed us to visit their collections and take data. Without their generous 
support, this study would not have been possible. 


LITERATURE CITED 


ALBRECHT, C. W., JR. 1982. The taxonomy, geography, and seasonal distribution of 
Rhopalocera in Ohio. Ph.D. Thesis, The Ohio State Univ. 512 pp. 

ANONYMOUS. 1984. Endangered and threatened wildlife and plants; review of inver- 
tebrate wildlife for listing as endangered and threatened species. U.S. Fish and 
Wildlife Service, Notice of Review. Federal Register 49:21664—21675. 

BALES, B. R. 1909. A partial list of the Lepidoptera of Pickaway County, Ohio. Entomol. 
News 20:169-177. 

BARNES, W. & J. MCDUNNOUGH. 1918. Illustrations of the North American species of 
the genus Catocala. Mem. Am. Mus. Nat. Hist. New Series Vol. 3, Part 1. 47 pp. 

BuBna, M. 1897. Entomology at Cleveland, Ohio. Entomol. News 8:97-99. 

COOPERRIDER, T. S. 1982. Endangered and threatened plants of Ohio. Ohio Biol. Surv., 
Biol. Notes No. 16. 92 pp. 

Cusick, A. W. & G. M. SILBERHORN. 1977. The vascular plants of unglaciated Ohio. 
Bull. Ohio Biol. Surv., New Series Vol. 5, No. 4. 157 pp. 

Diric, R. & J. F. CRYAN. 1976. The Karner blue project: January 1973 to December 
1976. Atala 4:22-26. 

Dury, C. 1876. List of Catocalae observed in the vicinity of Cincinnati, Ohio, 1876. 
Can. Entomol. 8:187-188. 

1878. Catalogue of the Lepidoptera observed in the vicinity of Cincinnati, Ohio, 
including diurnals, Sphingidae, Aegeridae, Zygaenidae, Bombycidae, Noctuidae, Pha- 
laenidae, and Pyralidae. J. Cincinnati Soc. Nat. Hist. 1:12-23. 

FORBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Part III. 
Noctuidae. Cornell Univ. Agr. Exp. Stat. Mem. 329. 433 pp. 

1960. Lepidoptera of New York and neighboring states. Part [V. Agaristidae 
through Nymphalidae, including butterflies. Cornell Univ. Agr. Exp. Stat. Mem. 371. 
188 pp. 

GALL, L. F. 1984. The evolutionary ecology of a species-rich sympatric array of Catocala 
moths. Ph.D. Thesis, Yale Univ. 264 pp. 

HAMMOND, P. C. & D. V. MCCORKLE. 1983. The decline and extinction of Speyeria 








12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


populations resulting from human environmental disturbances (Nymphalidae: Ar- 
gynninae). J. Res. Lepid. 22:217-224. 

HENNINGER, W. F. 1910. The macro-Lepidoptera of Seneca County, Ohio. Ohio Nat. 
11:233-242. 

HINE, J. S. 1898a. List of butterflies known to have been taken in Ohio. Ohio Acad. 
Sci., 6th Ann. Rpt. Pp. 22-27. 

1898b. Ohio butterflies. J. Proc. Columbus Hort. Soc. 12:77-79. 

HOLLAND, W. J. 1903. The moth book. Doubleday, Page & Co., New York. 479 pp. 

1931. The butterfly book. Doubleday, Garden City, New Jersey. 424 pp. 

KING, C. C. 1979. Hill country, pp. 161-181. In Lafferty, M. B. (ed.), Ohio’s natural 
heritage. Ohio Acadamy of Science, Columbus, Ohio. 

KIRTLAND, J. P. 1854. Diurnal Lepidoptera of northern and middle Ohio. Proc. Cleve- 
land Acad. Nat. Sci. 2:5, 73-75. 

Macy, R. W., & H. H. SHEPARD. 1941. Butterflies, a handbook of the butterflies of the 
United States, complete for the region north of the Potomac and Ohio rivers and 
east of the Dakotas. Univ. Minnesota Press, Minneapolis. 247 pp. 

MCALPINE, W. S., S. P. HUBBELL & T. E. PLISKE. 1960. The distribution, habits, and 
life history of Euptychia mitchellii (Satyridae). J. Lepid. Soc. 14:209-226. 

MILLER, W. E. 1979. Fire as an insect management tool. Bull. Entomol. Soc. Amer. 25: 
137-140. 

NaBokov, V. 1949. The nearctic forms of Lycaeides Hubner. (Lycaenidae, Lepidop- 
tera). Psyche 50:87-99. 

OPLER, P. A. & G. O. KRIZEK. 1984. Butterflies east of the Great Plains. Johns Hopkins, 
Baltimore, Maryland. 294 pp. 

PALLISTER, J. C. 1927. Cissia mitchelli (French) found in Ohio, with notes on its habits. 
Lepidoptera-Satyridae. Ohio J. Sci. 27:203-204. 

PILATE, G. R. 1882. List of Lepidoptera taken in and around Dayton, Ohio. Papilio 2: 
65-71. 

Price, H. F. 1970. Butterflies of northwestern Ohio. Mid-Continent Lepid. Ser., No. 
14, 16 pp. 

PRINGLE, J. S. 1980. An introduction to wetland classification in the Great Lakes region. 
Royal Bot. Gardens. Tech. Bull. No. 10. 11 pp. 

RAWSON, G. W. & J. S. THomas. 1939. The occurrence of Hemiargus isola (Reakirt) 
in northern Ohio. Bull. Brooklyn Entomol. Soc. 34:9-10. 

RUTKOWSKI, F. 1966. Rediscovery of Euptychia mitchellii (Satyridae) in New Jersey. 
J. Lepid. Soc. 20:43-44. 

SARGENT, T. D. 1976. Legion of night: The underwing moths. Univ. Massachusetts 
Press, Amherst. 222 pp. 

SCHWEITZER, D. F. 1982. The larva and status of Catocala pretiosa (Noctuidae), with 
designation of a lectotype. J. Lepid. Soc. 36:18-30. 

SHUEY, J. A. 1985. Habitat associations of wetland butterflies near the glacial maxima 
in Ohio, Indiana and Michigan. J. Res. Lepid. 24:176-186. 

STUCKEY, R. & G. DENNY. 1981. Prairie fens and bog fens in Ohio: Floristic similarities, 
differences, and geographical affinities. In Romans, R. C. (ed.), Geobotany II. Plenum, 
New York. 271 pp. 

STUDEBAKER, O. & D. STUDEBAKER. 1967. Butterflies of Miami County, Ohio. Newsletter 
Assoc. Minnesota Entomol. 1:61-65. 

Wyss, A. 1930. A list of Rhopalocera taken in Cincinnati and vicinity. Proc. Junior Soc. 
Nat. Sci. 1(8&9):4-6. 

1932. Guide to the butterflies of Cincinnati and vicinity. Junior Soc. Nat. Sci. 

Misc. Publ., No. 7. 37 unpaged. 











Received for publication 23 May 1986; accepted 14 November 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 18-22 


MILKWEED PATCH QUALITY, ADULT POPULATION 
STRUCTURE, AND EGG LAYING IN THE 
MONARCH BUTTERFLY 


M. P. ZALUCKI AND Y. SUZUKE 


Department of Entomology, University of Queensland, 
St. Lucia, Queensland 4067, Australia 


ABSTRACT. Relations between hostplant patch attributes (patch size, plant density, 
plant age, nectar availability), adult population structure (population size, sex ratio, age 
structure), and measures of egg laying (number of eggs per plant, total number of eggs 
per patch) in Danaus plexippus (L.) and its hostplant Asclepias fruticosa L. were inves- 
tigated. Sex ratios were male biased in areas with high hostplant density (patches), and 
female biased in nonpatch areas. Numbers of eggs per plant were higher on single, isolated 
plants (areas of low hostplant density) than on patch plants. Relations between adult 
population attributes and measures of egg laying were not clear-cut. Contrary to expec- 
tations, neither nectar availability nor sex ratio influenced measures of egg laying. Patch 
size and number of females in a patch were also unrelated to total number of eggs in a 
patch. However the last was positively related to number of males and percentage of 
young females in a patch, and negatively correlated with percentage of young males in 
a patch. 

Additional key words: Danaus plexippus, Nymphalidae, Asclepias fruticosa, Aus- 
tralia. 


Boggs and Gilbert (1979) demonstrated that mating provides not only 
sperm but also nitrogen-rich nutrients for egg production by females 
in the monarch butterfly, Danaus plexippus (L.), and two heliconid 
species. This nutritional contribution may be most important in multi- 
mating species which produce large spermatophores, such as D. plex- 
ippus (Burns 1968, Pliske 1978, Suzuki & Zalucki 1986). Bull et al. 
(1985) and Suzuki and Zalucki (1986) showed experimentally that fe- 
male D. plexippus are more likely to remain in and around patches of 
the milkweed Asclepias fruticosa (L.) where the sex ratio [males/(males 
and females)| is greater than 0.5. This implies that males are a major 
limiting resource for females. 

Many factors in addition to males influence egg laying on hostplants, 
including species, size, age, and condition of the plants (Zalucki & 
Kitching 1982a). Egg laying in patches of plants is further influenced 
by patch size, rules of female movement (Zalucki & Kitching 1982a, 
1982b, Zalucki 1983), and female age (Zalucki 1982). In this study, we 
looked for relations among monarch population attributes around patches 
(population size, sex ratio, age, and distribution), patch characteristics 
(size, plant density and age, nectar availability) and egg “investment” 
(number of eggs per plant, total number of eggs per patch), after 


‘ Present address: Department of Biophysics, Kyoto University, Kyoto 606, Japan. 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Effect of patch category on sex ratio (proportion male) in two sampling 
periods. 


Sampling period 


24-31 Aug. 1983 20 Feb.—1 Mar. 1984 
Patch No. of Mean + SD Range of No. of Mean + SD Range of 
- category patches sex ratio sex ratios patches sex ratio sex ratios 
Nonpatch 4 0.38 + 0.11 0.24—0.50 4° 0.43 + 0.03 0.40-0.47 
Small 8 0.54 + 0.10 0.35—0.66 7 0.61 + 0.09 0.43—0.69 
Medium 1 0.63 = 4 0.65 + 0.07 0.58-0.72 
Large 2 0.46 + 0.01 0.45—0.47 2 0.61 + 0.08 0.55—0.67 


2 Excludes 1 misclassified nonpatch. 


recording these attributes from a large and diverse sample of milkweed 
patches over a short duration. 


MATERIALS AND METHODS 


Milkweed patches were sampled on two occasions giving a total of 
33 patches. During the first sampling period (24 to 31 August 1983), 
monarchs in and very near 15 patches were netted for 1 person h at 
each patch. All butterflies caught were sexed and scored for age on the 
basis of wing condition as follows: A’—wing soft, A—wings intact, B— 
wings frayed, C—wings very frayed. In this first survey, patches were 
classified subjectively: nonpatch (milkweed scattered and at very low 
density, <1 plant/m?), small patch, medium, or large patch. Egg num- 
bers were not recorded. This survey could be used only to ascertain 
relations between patch category and butterfly sex ratio. Patches were 
located in and near Beenleigh (27°43’S, 153°12’E), Logan Village (27°46’S, 
153°06’E), and Mt. Crosby (27°32’S, 152°42’E) in SE Queensland, Aus- 
tralia. 

On the second sampling occasion (20 February to 1 March 1984), 18 
patches located along the Kenmore (27°30'S, 152°56’E)—Mt. Crosby 
Road were used. Adults were netted for 1 person h per patch, sexed, 
and aged as before. An attempt was made to estimate adult sampling 
efficiency. In smaller patches it is easier to obtain a high sampling 
fraction than in large patches. Sampling fractions were estimated sub- 
jectively on the basis of butterflies seen after netting ceased. Sampling 
fraction is the ratio of butterflies caught in 1 person h to butterflies 
caught plus those seen flying at the end of netting. Patch size was 
measured by idealizing the patch to standard shapes (circle, square, 
etc.) and recording relevant distances using a tape measure. Plant den- 
sity within a patch was estimated using 6 to 25 randomly thrown 1-m? 
quadrants. From each patch a sample of ca. 100 plants was selected, 
cut at ground level, placed in plastic bags, and returned to the labo- 


VOLUME 41, NUMBER 1 15 


TABLE 2. Details of sampling during 20 February—1 March 1984. 


es time Temperature Patch Sampling 
( 


Date range (°C) Remarks category* Sex ratio fraction 

20-2-84 0945 21-29 Cloudy, 3 0.69 0.80 
1045 showers 2 0.69 0.80 

21-2-84 0855 22-30 Cloudy, 2 0.66 0.70 
0950 fine ] 0.47 0.30 

1040 3 0.72 0.70 

1145 2 0.69 1.00 

22-2-84 0910 22-27 Cloudy 2, 0.56 0.70 
1000 4 0.55 0.30 

1100 3 0.61 0.80 

23-2-84 0910 21-29 Cloudy, 3 0.58 0.60 
1000 showers 2 0.43 0.30 

1055 4 0.67 0.70 

28-2-84 0935 22-36 Fine 2 0.67 0.30 
1015 1 0.43 0.80 

1100 2 0.67 0.60 

1-3-84 0900 20-27 Cloudy 2 0.59 1.00 
0945 ] 0.44 0.70 

1035 1 0.43 0.80 


i 1 = nonpatch; 2 = small patch, <20,000 plants; 3 = medium patch, 20-80,000 plants; 4 = large patch, >80,000 
plants. 


ratory. These were measured and classified as young, with flowers, with 
small pods, with old pods, or dead. They were then searched for eggs 
and larvae. Percentage of flowering milkweed plants provided an es- 
timate of nectar availability. Other potential nectar resources (lantana, 
weeds) were scored subjectively as low, medium, or high. 


RESULTS 


Data from both sampling periods were combined to look for relations 
between patch category and sex ratio (Table 1). A two-way ANOVA 
on an arcsin (square root) transformation (Sokal & Rohlf 1981) indicated 
highly significant effects of patch category (P < 0.001) and sampling 
period (P < 0.05). However, removing the nonpatch category from 
analyses removed the effects of both patch category (P > 0.05) and 
sampling period (P > 0.05) on sex ratio. Nonpatches had consistently 
female-biased sex ratios (Table 1) whereas patches, regardless of size, 
had variable but generally male-biased sex ratios (Table 1). 

More detailed observations were made during the second sampling 
period. Neither time of day nor weather had any obvious effect on sex 
ratios or sampling fractions (Table 2). Sampling fraction is a function 
of patch size and terrain. Smaller sampling fractions were recorded for 
large patches and for patches on hillsides or with long grass. 

Relations between various measures for eggs laid, patch, and butterfly 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


16 


‘(Que[d /s33a ‘ou ueeul) x (sjuejd ‘ou [e}07) » 
‘wuoKoeIy Suldures 10} pa}d9110F , 
‘(,u1/syuejd ou) x (,Ur ‘azIs yo}eg)- 


SSS SaaS Ne NG 


¢9'0-— €L0O = +89°0 *VL0 880 cs'0 PIX 83839 ‘OU [B]0], 

*LG 0 ClX & sunok ¥ 

6GO- IX 9 Bunok % 

*09 O0— IlX O1}e1 Xa 
160 860 cSO- 90 *68'0 OS 0 89°0 OLX ‘OU [2}0], 

060 1g 0 *V80 670 S90 6X qo “ON 

SD SO) 060 £8sF0 L9°0 8x q? ON 

ec'o— LX queld /s339 ‘oN 

0c ‘0 OX ase Ur g 

CX Iejeu 19410 

PX BULIOMOT % 

L8°0 SX  Sjueyd ‘ou [e}0], 

ox Ayisuap jury g 

IX eZIs yoeg 








VIX SIX OLX 11x OLX 6X 8X LX 9X OK vx SX GX IX IESMLS/ Ree 


‘syuIod SuIA]}NO OM} IO BUO 0} enp 
QOULOFIUBIS SOJOIPUI y “‘UMOYS d1e CO':N > ge JULOFIUSIS syUsIOyJa00 AJUO ‘sayoyed ET Ul SayqerieA |e OJ XIIQeUr uOHneeIIOD =-“§ ATAV] 


VOLUME 41, NUMBER 1 1 67/ 


Eggs/plant (x 100) 
90 


80 e 


2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 


Plant density/m 2 


Fic. 1. Effect of plant density on number of eggs per 100 plants. Points with plant 
density 0 correspond to nonpatches. 


variables were compared using all possible pairwise correlations, ig- 
noring nonpatches. The correlation matrix (Table 3) shows only coef- 
ficients significant at P < 0.05 (one-tailed t-test). Some correlations 
were “significant” due to one or two outlying points. These relations 
are probably superficial. 

A number of patterns are apparent in Table 3. Patch-size variables 
(area and total number of plants in patch) and numbers of adult but- 
terflies were positively correlated with total numbers of eggs in patches. 
However, numbers of eggs per plant were negatively related to plant 
density (Fig. 1) and patch size. Plant age (% plants with pods and dead 
stems) correlated positively with numbers of butterflies, but this may 
follow from the positive association between total numbers of plants 
and plant age. 

One striking result was that neither number of females nor percentage 
of young females in a sample had any influence on total number of 
eggs or number of eggs per plant, respectively, once outlying points 
were removed. Equations for these relations are: Total no. eggs (y) = 
167 x No. females (x) + 1,775, F,,, = 9.498, P < 0.05 when all points 
included; deleting one outlier, y = 25x + 4,177, F, 1) = 0.079, P > 0.05; 
and No. eggs/plant (y) = 0.57 x % young females (x) — 1.87, F\,, = 
5.277, P < 0.05 when all points included; deleting two outliers, y 
0.091x + 10.16, F,, = 0.144, P > 0.05). 


18 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TOTAL EGGS (x 100) 





O 
0 10 20 30 40 50 60 70 80 90 100 110 
MALES IN SAMPLE 


Fic. 2. Effect of number of males caught on total number of eggs in a patch. 


On the other hand, number of males in a sample had a negative 
association with number of eggs per plant, a positive one with total 
number of eggs (Fig. 2), and the latter was negatively related to per- 
centage of young males in a patch (Fig. 3). The positive association 
between number of males (x) and total number of eggs (y) was signif- 
icant (P < 0.05) only if all points were included (Fig. 2, y = 158x + 
37.8, F,,, = 18.62). Deleting the extreme value from this figure removes 
significance (y = 67.39x + 2,585, F,,) = 0.8958, P > 0.3). This was not 
the case for the relation between total number of eggs (y) and % young 
males (x) (Fig. 3). Removing one or two extreme values did not change 
this relation (with the two extreme left-hand points deleted, y = 
—295.7x + 17,676, F,, = 5.728, P < 0.05). The negative association of 
number of eggs per plant and number of males probably stems from 
the negative relations of number of eggs per plant to total plant number 
and density, and the strong positive association between these variables 
and male number (Table 3). Neither nectar availability nor sex ratio 
influenced adult numbers or measures of egg laying. 

For any set of data where most variables are correlated and inter- 
dependent, single pairwise comparisons may be misleading. From the 
above analysis the variables total number of eggs in a patch, patch size, 
numbers of males and females, and percentages of young males and 


VOLUME 41, NUMBER 1 19 


TOTAL EGGS (x100) 
200 


\ Y =22988-398x 
120 
4 F(1,!l) =8.153,P < 0.02 


e \ 
80 \te ® 
o\ 





10 20 30 40 50 60 70 80 90 100 
% YOUNG MALES 


Fic. 3. Relation of percentage of young males to total number of eggs in a patch. 


females in a patch were entered in a partial correlation matrix (Table 
4). Total number of eggs in a patch was positively correlated with male 
density, but negatively correlated with percentage of young males in 
the population. Total number of eggs laid and percentage of young 
females were also positively related (P < 0.10), as was egg number and 
percentages of young males and females (P < 0.05). Surprisingly neither 
patch size nor number of females was related to total number of eggs 
laid. 

These analyses were corroborated by stepwise linear regression with 
total number of eggs as the dependent variable. Results (Table 5) in- 
dicated significant effects only for number of males, % young males, 
and % young females. 


DISCUSSION 


Zalucki and Kitching (1985) and Bull et al. (1985) found that sex 
ratios tend to be male-biased in and around milkweed patches. If, as 
they suggest, male bias cannot be explained by sex ratio at birth, survival 
difference, or sampling bias within a patch, then where do the females 
go? The present study confirms male bias around milkweed patches 
(ca. 60%) and demonstrates a female-biased population outside patches 
(nonpatch areas), hilltops notwithstanding. As with many Lepidoptera 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Partial correlation matrix for selected patch, butterfly, and egg-laying vari- 
ables. 


Number % young 
Total eggs Patch size Males Females Males Females 
X1 X2 X38 X4 X5 X6 
x2 0.52 
x3 O24" =()) IL 7/ 
x4 —0.37 0.24 0.80** 
xO —0.79** 0.13 0.69** —0.40 
x6 0.62* —0.10 —0:52 0.20 0.60* 


**P < 0.01; *P < 0.05. 


(Shields 1967, Parker 1978, others) male monarchs can be taken in 
numbers around hilltops. 

The egg-laying pattern found by Zalucki and Kitching (1982a) on 
artificial milkweed “‘patches” was also confirmed by this study. Number 
of eggs per plant was much higher on single isolated plants than on 
patch plants (Fig. 1). This result can be explained in part by how female 
monarchs search for and use a host plant (Zalucki & Kitching 1982b, 
Zalucki 1983), and by female-biased sex ratios outside patches (single 
plants). Jones (1977) found a similar pattern of egg-laying in Pieris 
rapae (also see Shapiro 1981, Thompson & Price 1977, Wiklund & 
Ahrberg 1978). However, there are many alternate explanations for 
such a pattern (Mackay & Singer 1982). 

Within patches, reiations among patch and butterfly variables and 
measures of egg-laying are not straightforward. Contrary to expectation, 
sex ratio was not related to any measure of egg-laying. Nor was number 
of eggs in a patch related to availability of nectar, number of females 
in a patch, or patch size (Tables 3-5). Bull et al. (1985) and Suzuki and 
Zalucki (1986) showed experimentally that female residence time in a 
patch is positively related to sex ratio, and Bull et al. (1985) could not 
find any effect of butterfly density on residence time. Neither study 
recorded egg numbers in experimental patches. 

In the present study, number of eggs in a patch was positively related 
to number of males (all ages), but not to number of females, even 
though the latter two variables are strongly positively associated (Tables 
3, 4). This provides further circumstantial evidence that males are a 
major “egg-laying resource” for females through provision of nutrients 
as well as sperm at mating (Boggs & Gilbert 1979, Suzuki and Zalucki 
1986). Also, Herman and Barker (1977) showed that mating stimulates 
oogenesis. There was a weak positive relation between female age 
structure and number of eggs; patches with a high percentage of young 
females had more eggs. This presumably reflects the high fecundity of 
such females (Zalucki 1982). 


VOLUME 41, NUMBER 1 91 


TABLE 5. Stepwise linear regression of total number of eggs against patch size, number 
of males and females, and % young males and females. 


Independent Coefficient Standardized 
variable (+SE) Coefficient t-value P 
Patch size 18.88 (11.67) 0.28 1.618 NS 
No. males 153.40 (55.54) 0.72 ZOOL, ae 
No. females —65.18 (61.35) —0.27 —1.062 NS 
% young males — 271.40 (80.34) —0.44 —3.379 og 
% young females 69.16 (33.21) 0.23 2.083 if 


**P < 0.05; *P < 0.10. 


In contrast, number of eggs in a patch was inversely related to per- 
centage of young males in the population. Are young males more 
aggressive (inexperienced) in courtship and do they subsequently “drive” 
females from a patch? This hypothesis will require further testing. 


ACKNOWLEDGMENTS 


We thank R. L. Kitching for reading an earlier version of this manuscript, T. Barnes 
for statistical advice, and B. Dennis for typing. The junior author was supported by a 
University of Queensland Postdoctoral Fellowship. 


LITERATURE CITED 


Bocecs, C. & L. GILBERT. 1979. Male contribution to egg production in butterflies: 
Evidence for transfer of nutrients at mating. Science 206:83-84. 

Bunmeeo Me M. Pe ZALUCKI, Y. SUZUKI, D. A. MACKAY & R. L. KITGHING. 1985. An 
experimental investigation of resource use by female monarch butterflies, Danaus 
plexippus (L.). Aust. J. Ecol. 10:391-398. 

BuRNS, J. M. 1968. Mating frequency in natural populations of skippers and butterflies 
as determined by spermatophore counts. Proc. Nat. Acad. Sci. U.S.A. 61:852-859. 

HERMAN, W.S. & J. F. BARKER. 1977. Effect of mating on monarch oogenesis. Experien- 
tia 15:688-689. 

JONEs, R. E. 1977. Movement patterns and the egg distributions of cabbage butterflies. 
J. Anim. Ecol. 46:115-1235. 

Mackay, D. A. & M. C. SINGER. 1982. The basis of an apparent preference for isolated 
host plants by ovipositing Euptychia libye butterflies. Ecol. Entomol. 7:299-303. 
PARKER, G. A. 1978. Evolution of competitive mate searching. Ann. Rev. Entomol. 23: 

173-196. 

PLISKE, T. E. 1973. Factors determining mating frequency in some New World but- 
terflies and skippers. Ann. Entomol. Soc. Am. 66:164-169. 

SHAPIRO, A. M. 1981. The pierid red-egg syndrome. Am. Natur. 117:276-294. 

SHIELDS, O. 1967. Hilltopping: An ecological study of summit congregation behaviour 
of butterflies on a southern California hill. J. Res. Lepid. 6:69-178. 

SOKAL, R. R. & F. J. ROHLF. 1981. Biometry. 2nd ed. W. H. Freeman, San Francisco. 
859 pp. 

SuzuUKI, Y. & M. P. ZALUCKI. 1986. Mate acquisition as a factor influencing female 
dispersal in Danaus plexippus (L.) (Lepidoptera: Danaidae). Aust. J. Entomol. Soc. 
25:31-35. 

THOMPSON, J. N. & P. W. Price. 1977. Plant plasticity, phenology and herbivore 
dispersion: Wild parsnip and the parsnip webworm. Ecology 58:1112-1119. 

WIKLUND, C. & C. AHRBERG. 1978. Host plants, nectar source plants and habitat 
selection of males and females of Anthocharis cardamines (Lepidoptera). Oikos 31: 
169-183. 


29 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ZALUCKI, M. P. 1982. The effects of age and weather on egg laying in Danaus plexippus 
L. (Lepidoptera: Danaidae). Res. Pop. Ecol. 23:318-327. 

1983. Simulation of movement and egg laying in Danaus plexippus (Lepidop- 
tera: Nymphalidae). Res. Pop. Ecol. 25:353-365. 

ZALUCKI, M. P. & R. L. KITCHING. 1982a. The dynamics of oviposition of Danaus 
plexippus L. on Asclepias spp. J. Zool. Lond. 198:103-116. 

1982b. Movement patterns in Danaus plexippus L. Behaviour 80:174-198. 

1985. The dynamics of adult Danaus plexippus L. within patches of its food 

plant, Asclepias spp. J. Lepid. Soc. 38:209-219. 











ANNOUNCEMENT 
TECHNICAL COMMENTS: A NEW CATEGORY IN THE JOURNAL 


Readers are invited to submit Technical Comments up to 1,600 words on any research 
article on Lepidoptera published in any refereed journal within the previous two years. 
Technical Comments may offer additional information, alternative interpretations, or 
otherwise elaborate or footnote a published paper in some useful way. 

Authors of the original papers will be asked for their opinion of comments, and be 
given an opportunity to reply in the same Journal issue if the comments are accepted. 
Technical Comments and replies will be subject to the usual review procedures, and the 
former to page charges. Comments falling outside the above guidelines will be evaluated 
as General Notes or Articles. 

This new category is meant to provide a timely lepidopterology forum and thereby to 
increase the value of the Journal to readers and authors. 


WILLIAM E. MILLER, Editor 


Journal of the Lepidopterists’ Society 
41(1), 1987, 23-28 


HOST SPECIFICITY AND BIOLOGY OF 
BUCCULATRIX IVELLA BUSCK, A POTENTIAL 
BIOLOGICAL CONTROL AGENT FOR 
BACCHARIS HALIMIFOLIA L. IN AUSTRALIA 


W. A. PALMER 


North American Field Station, Queensland Department of Lands, 
2714 Pecan Drive, Temple, Texas 76502 


AND 


G. DIATLOFF 


Alan Fletcher Research Station, Queensland Department of Lands, 
27 Magazine Street, Sherwood, Queensland 4075, Australia 


ABSTRACT. Life history and host range of Bucculatrix ivella were investigated as 
part of a program to find host-specific biocontrol agents for Baccharis halimifolia in 
Australia. This multivoltine insect was collected on B. halimifolia and B. neglecta from 
Texas to New Jersey. Though usually found at low population densities, it occasionally 
occurred in numbers sufficient to defoliate plants. Host specificity tests of oviposition 
preference and larval feeding indicated that B. ivella was specific to Baccharis species. 
These tests and a field survey indicated that Iva frutescens is definitely not a host although 
it is reported as such in the literature. Bucculatrix ivella has been approved for introduction 
into Australia for the control of B. halimifolia. 


Additional key words: Lyonetiidae, Baccharis neglecta, Iva frutescens, introduction, 
weed. 


Following its introduction into Queensland, Australia, before 1900, 
the North American shrub Baccharis halimifolia L. (Asteraceae: As- 
tereae: Baccharineae) has become a serious weed in SE Queensland and 
NE New South Wales by invading pastures and land cleared for re- 
forestation. The plant was declared noxious in 1951; subsequently a 
biological control program to find and introduce suitable host specific 
insects from the New World was implemented. This program consisted 
of intensively surveying appropriate areas, selecting stenophagous species 
from available knowledge, testing the host range of these species ex- 
perimentally and, if their host range was limited to Baccharis, mass 
rearing and releasing in Australia. 

One such candidate from the surveys of the fauna on B. halimifolia 
and B. neglecta Britt. (Palmer 1987, Palmer & Bennett, in prep.) was 
the leafminer Bucculatrix ivella Busck. This insect was first reported 
on Iva frutescens L. (Busck 1904), but since has been reported only 
from B. halimifolia (Braun 1968, R. W. Hodges, pers. comm.). How- 
ever, we suspected the host record on I. frutescens may have been in 
error when another species, Aristotelia ivella Busck (Gelechiidae), col- 
lected by H. Dyar from the same plants (Busck 1904) proved to be 
specific to Baccharis. 


24 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The genus Bucculatrix is cosmopolitan with about half the species 
found in America north of Mexico. Braun (1963) described 99 species, 
including 50 new species, that occur in this region. Little taxonomic 
work has been done on the genus since, and there are undoubtedly 
many undescribed species (R. W. Hodges, pers. comm.). Although the 
known food plants include 25 plant families, nearly % of Bucculatrix 
species were associated with Asteraceae, and more than % were recorded 
from a single plant genus (Braun 1963). Braun (1963) listed three pest 
species: B. thurberiella Busck, B. canadensisella Chambers, and B. 
pomifoliiella Clemens, which attack cotton, birch, and apple, respec- 
tively. An undescribed Bucculatrix has already been released in Aus- 
tralia for control of the weed Parthenium hysterophorus L. 

At least four species are associated with Baccharis. In addition to 
Bucculatrix ivella, B. separabilis Braun, and B. variabilis Braun are 
associated with Baccharis pilularis DC. in California (Braun 1963). 
Recently, we collected an undescribed species from B. sarathroides 
Gray in Arizona. 

This paper reports results of experimental and field observations 
undertaken to investigate the host specificity of Bucculatrix ivella. In 
the course of the study, biology and phenology of B. ivella were observed 
and are also reported. 


BIOLOGY 


The biology of B. ivella is typical of leaf-mining species of Bucculatrix 
described by Braun (1963). The following description is based on nu- 
merous laboratory and field observations of life stages. Eggs were whit- 
ish, translucent, and flatish ovoid in shape, and were cemented to the 
leaf surface, usually along the upper surface of the midrib of the leaf. 
In sunlight they had a characteristic iridescence and were thus easily 
recognized. 

Larvae hatched within 7-11 days and entered the leaf tissue directly 
where they mined the parenchyma. Mines were distinctive threadlike 
tracks which initially followed a leaf vein but eventually became ir- 
regular, serpentine, and black with the deposition of frass. Larvae spent 
the first, second, and part of the third instars in these mines. After 
leaving the mines, they spun flat, thin “molting cocoons” under which 
they molted. Fourth and fifth instars were external feeders, and a second 
“molting cocoon’ was formed at the end of the fourth instar. Larvae 
were found on either upper or lower leaf surfaces, although the latter 
were more commonly infested. When disturbed, larvae dropped on a 
silken thread. They consumed leaf tissue in patches but left the opposite 
epidermal tissue intact, which produced a “window” effect. 


VOLUME 41, NUMBER 1 95 


TABLE 1. Number of plants infested with Bucculatrix ivella after exposure in a 
glasshouse. 


Plant species (Tribe) Number examined Number infested 


Baccharis halimifolia L. (Astereae) 8 
B. neglecta Britton (Astereae) 

B. glutinosa (R. & P.) (Astereae) 

B. pilularis DC. (Astereae) 

Solidago altissima L. (Astereae) 

Haplopappus tenuisectus (Green) Blake (Astereae) 
Aster novae-angliae L. (Astereae) 

Conyza canadensis (L.) (Astereae) 

Iva frutescens L. (Heleantheae) 

Leucanthemum maximum Ramond (Anthemideae) 
Ageratum houstonianum Mill. (Eupatoreae) 


— 
NOWOrFNNRFRE UU 
oooocooooeoonhnt 


Pupation occurred within characteristic ribbed cocoons spun on leaves 
and stems of Baccharis. However, when larvae were abundant, some 
left the host to pupate on neighboring plants or in ground debris. 

Before eclosion, the pupa thrust through the anterior end of the 
cocoon, exposing about half its length. Pupal cases remained attached 
to cocoons after moth emergence. Moths remained quiescent on foliage 
during the day and became active at dusk. 


Hosts, DISTRIBUTION AND PHENOLOGY 


Bucculatrix ivella has been collected nearly throughout the range of 
Baccharis halimifolia; we collected it in New Jersey, Virginia, Florida, 
Louisiana, and Texas. In Texas, the immatures were most abundant in 
late April and early May, when infested plants had 50-100 larval mines, 
but after early spring few individuals were seen. In Florida, the life 
cycle was completed in 4-6 weeks and larvae were found at most times 
of the year. Several hundred mines per plant were observed in Virginia 
and New Jersey in June and July, and these populations caused severe 
defoliation. It is perhaps the most abundant lepidopteran associated 
with B. halimifolia. 

In central Texas, Bucculatrix ivella was collected from Baccharis 
neglecta, which is a new host record. However, natural populations on 
this species were invariably low, even though it breeds readily on this 
plant under glasshouse conditions. 

Collections of Bucculatrix ivella immatures invariably contained a 
proportion (10-90%) of parasitized specimens. Among the parasites to 
emerge were species of the hymenopterans Ageniaspis (Encertidae), 
Apanteles (Braconidae), Bucculatriplex (Braconidae), Cirrospilus (Eu- 
lophidae), Mirax (Braconidae), Opius (Braconidae) and Tetrastichus 
(Eulophidae). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


bo 
fo») 


TABLE 2. Degree of infestation by Bucculatrix ivella on six plants in an unreplicated 
cage experiment. 





Plant species (Tribe) Infestation 





Baccharis halimifolia L. (Astereae) 13 larval mines 
20 larval cocoons 
5 late instars 
external feeding damage 
>60 pupal cocoons 


Aster novae-angliae L. (Astereae) none 
Solidago altissima L. (Astereae) none 
Callistephus chinensis (L.) Nees (Astereae) none 
Cynara scolymus L. (Cardueae) none 
Iva frutescens L. (Heleantheae) none 


HOST SPECIFICITY 


Host specificity was determined by laboratory trials and field observations designed 
first to demonstrate whether Iva frutescens in particular, and Asteraceous plants in general, 
were hosts, and second, to test the insect against a wide variety of plants of economic 
importance to Australia. 

Field observations. In Virginia and New Jersey, Baccharis halimifolia heavily infested 
with Bucculatrix ivella was found growing close to Iva frutescens. Sometimes the branches 
of the two intertwined. Iva frutescens was carefully searched without ever finding an 
infestation of Bucculatrix ivella. 

The area where the report on Iva frutescens originated (Palm Beach, Florida) was also 
searched. However, I. frutescens was not found there; it is apparently rare S of Daytona 
Beach, Florida, while Baccharis halimifolia is common. 

Larval feeding. On four occasions over a two-year period, late instars were collected 
from B. halimifolia and returned to the laboratory where 5 to 10 were placed on bouquets 
of B. halimifolia and I. frutescens. Larvae invariably continued feeding and pupated on 
the B. halimifolia, but no evidence of feeding was found on the I. frutescens. 

Glasshouse observations. Potted asteraceous plants were introduced into a glasshouse 
containing potted Baccharis plants infested with Bucculatrix ivella. After two months, 
the plants were carefully examined for infestations of B. ivella. Only Baccharis halimifolia 
and B. neglecta were infested (Table 1). 

Cage experiment. A cage experiment was conducted ina glasshouse using B. halimifolia 
and five other asteraceous species common in Australia. One potted plant of each species 
was placed in a wooden cage with a clear plastic top. Thirty pupae and four moths were 
introduced together with four sugar-water wicks. When the B. halimifolia plant was 
infested, all plants were removed and carefully examined. A number of infestations were 
found on the B. halimifolia, but none of the other plants was infested (Table 2). 

Comprehensive testing. Before permission to release the insect in Australia could be 
sought, Bucculatrix ivella had to be tested against the complete list of plants suggested 
by the Commonwealth Department of Health (Table 3). Oviposition preference was 
tested using a 5.0 x 4.5 x 3.0 m glass-sided cage into which were randomly placed young, 
actively growing tip cuttings of all test plants and Baccharis halimifolia. These cuttings 
were held in glass vials with water. Twenty moths were placed in the cage together with 
a honey-water mixture. After 6 days, when the B. halimifolia was infested with eggs, the 
cuttings were carefully examined and any eggs counted. After a further six days, when 
larval mines were seen in B. halimifolia leaves, the cuttings were reexamined for eggs 
or mines. The experiment was replicated twice. Numbers of eggs and larval mines were 
seen on all B. halimifolia cuttings, but not on any other cutting. 

_ In a second experiment, host range of the ectophagous late instars was tested. Five 
fourth or fifth instars collected from the field were placed on two potted plants of each 


VOLUME 41, NUMBER 1 97 


TABLES. Plant species against which Bucculatrix ivella was tested to obtain permission 
for its introduction into Australia. 


Apiaceae: Daucus carota L.; Pastinaca sativa L. 

Anacardiaceae: Mangifera indica L. 

Asteraceae: Baccharis halimifolia L.; Carthamus tinctorius L.; Chrysanthemum sp.; 
Dhalia sp.; Helianthus annuus L.; Lactuca sativa L. 

Brassicaceae: Brassica oleraceae (L.) Alef.; Brassica rapa L. 

Bromeliaceae: Ananas comosus (L.) Merr. 

Caricaceae: Carica papaya L. 

Chenopodiaceae: Beta vulgaris L. 

Convolvulaceae: Ipomoea batatas (L.) Lam. 

Cucurbitaceae: Cucumis melo L.; Cucumis sativus L.; Curcubita maxima Duch. 

Fabiaceae: Arachis hypogaea L.; Centrosema pubescens Benth.; Desmodium canum 
(Gmel.); Glycine wightii (R. Grah. ex Wight & Arn.) Verdc.; Glycine max (L.) Merr.; 
Medicago sativa L.; Phaseolus atropurpureus DC.; Phaseolus vulgaris L.; Pisum sa- 
tivum L.; Stizolobium sp.; Stylosanthes gracilis; Trifolium repens L.; Vigna catjang V. 

Linaceae: Linum usitatissimum L. 

Malvaceae: Gossypium hirsutum L. 

Mimosaceae: Leucaena leucocephala (Lam.) de Wit. 

Musaceae: Musa sapientum M. 

Passifloraceae: Passiflora edulis Sims 

Pinaceae: Pinus radiata D. Don.; Pinus taeda L. 

Poaceae: Avena sativa L.; Digitaria decumbens Stent.; Panicum maximum Jacq.; Pas- 
palum dilatatum Poir.; Pennisetum clandestinum Chiov.; Saccharum officinarum L.; 
Sorghum vulgare L.; Triticum aestivum L.; Zea mays L. 

Proteaceae: Macadamia integrifolia Maid & Betche 

Rosaceae: Fragaria vesca L.; Malus sylvestris Mill., Prunus domestica L.; Prunus persica 
(L.) Batch.; Pyrus communis L.; Rosa sp. 

Rutaceae: Citrus limon (L.) Burm. F.; Citrus paradisi Macfady.; Citrus reticulata Blanco; 
Citrus sinsensis (L.) 

Sapindaceae: Litchi chinensis Sonn. 

Solanaceae: Capsicum annuum L.; Lycopersicum esculentum Miller; Nicotiana tabacum 
L.; Solanum tuberosum L. 

Vitaceae: Vitis vinifera L. 

Zingiberaceae: Zingiber officinale Roscoe. 


species, which were observed for 10 days. Larvae on all plants other than B. halimifolia 
were seen leaving the plants in the first two days, and no feeding was attempted on them. 
On the other hand, all larvae on B. halimifolia were seen to feed normally. 


DISCUSSION 


The tests conducted on Bucculatrix ivella clearly showed its speci- 
ficity to some species of Baccharis such as B. halimifolia and B. neglecta; 
also that Iva frutescens was not a suitable host. The most likely expla- 
nation for the record on I. frutescens is that B. halimifolia was misi- 
dentified as that species. The two are morphologically similar and grow 
in similar habitats. They are both known by the same common name, 
“salt bush’. A second explanation is that if the two plant species were 
growing together, late instars could have migrated onto the I. frutescens 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


to pupate. This is less likely because I. frutescens does not grow, or is 
rare, in Palm Beach Co., Florida, where the collection was made. 

Bucculatrix ivella has considerable potential as a biocontrol agent 
and is also relatively easy to rear and handle in the laboratory. It is 
multivoltine and capable of building up to high populations that greatly 
damage the plant. In North America, it is heavily parasitized, but before 
release in Australia it will be freed of parasites. In the absence of 
parasites, the rate of population growth in Australia should be greater 
than in its native habitat. 

Permission to import this insect into Australia was granted in 1986. 


ACKNOWLEDGMENTS 


We thank D. R. Davis of the National Museum of Natural History, Smithsonian Institute, 
and R. W. Hodges of the Systematic Entomology Laboratory, USDA, who provided expert 
identification of collected material, and advised on some aspects of the study. 


LITERATURE CITED 


BRAUN, A. F. 1968. The genus Bucculatrix in America north of Mexico (Microlepidop- 
tera). Mem. Am. Entomol. Soc. 18:1—208. 

Busck, A. 1904. New species of moths of the superfamily Tineina from Florida. Proc. 
U.S. Natl. Mus. 28:225. 

PALMER, W. A. 1987. The phytophagous insect fauna associated with Baccharis ha- 
limifolia L. and B. neglecta Britton in Texas, Louisiana and northern Mexico. Proc. 
Entomol. Soc. Wash. 89:185-199. 


Received for publication 14 October 1986; accepted 8 December 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 29-40 


BUTTERFLIES FROM THE UJUNG KULON 
NATIONAL PARK, INDONESIA 


T. R. NEw! 


Department of Zoology, La Trobe University, 
Bundoora, Victoria 3083, Australia? 


M. B. BUSH 


Department of Geography, University of Hull, 
Humberside, United Kingdom 


AND 


H. K. SUDARMAN 


Museum Zoologicum Bogoriensis, Bogor, Indonesia 


ABSTRACT. Butterflies found on the Ujung Kulon Peninsula (85 spp.) and Pulau 
Peucang (36 spp.) in 1984 are listed and compared with the fauna of the Krakatau Islands. 
Many species are limited to particular vegetation types or ecotones, and maintenance of 
habitat diversity appears vital to ensure continued diversity in the national park. At 
present, management for grazing by large mammals and limited human access are 
practical conservation measures for butterflies. 


Additional key words: Java, surveys, conservation, biogeography. 


The Ujung Kulon Peninsula and adjacent islands comprise the west- 
ernmost region of Java, Indonesia. Since the large-scale inundations by 
tsunamis associated with the eruption of Krakatau in 1888, this region 
has been largely free from human activities. It is now a national park 
to which access is strictly controlled and monitored, and in which 
scientific work may be undertaken only by permit. Land access is 
difficult, and most visitors arrive by boat from towns on the NW coast 
of Java and stay predominantly on Pulau Peucang (Fig. 1). The main 
purpose of this park, which comprises some 30,000 ha, is to foster the 
last remaining population of the Javan rhinoceros, Rhinoceros sondaicus 
sondaicus Desmarest, but the relatively undisturbed forest and other 
vegetation types render it one of the most significant areas in Java for 
the conservation of native fauna. Much of the forest survived the 1883 
catastrophe, but little forest now present is considered to be true mature 
lowland rainforest (Blower & van der Zon 1977). The latter occurs on 
Gunung Payung, the highest point of the western part of the park, and 
on the nearby island of Pulau Peucang (Fig. 1). Much of the rest of 
western Java has undergone considerable change in support of a bur- 
geoning human population, and there is little doubt that the Ujung 
Kulon National Park now harbors remnant populations of many taxa 


' Address for correspondence. 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


which were formerly much more widespread. In common with much 
of the “third world’’, the distribution of reserves tends to reflect human 
settlement patterns, but the significance of Ujung Kulon was recognized 
by bestowal of the status of ‘nature reserve” in 1921 (Hoogerwerf 1970). 
Since that time it has been upgraded to a national park, and Ujung 
Kulon has become “the most widely known conservation area in South 
East Asia” (Blower & van der Zon 1977). It assumes additional biogeo- 
graphical importance as a likely major source area of animals and plants 
that have reached the Krakatau Islands over the last century since life 
there was expunged in 1883. 

In 1984, La Trobe University and the Bogor Zoological Museum 
mounted a joint expedition to survey the fauna of the Krakatau Islands, 
some 60 km N of the Ujung Kulon Peninsula, but politically part of 
the same national park. Nearly 30 biologists participated in the ex- 
pedition (Thornton 1985, Thornton & Rosengren 1987). An auxiliary 
aim was to examine the biota of selected areas bordering the Sunda 
Strait to assess likely origins of the Krakatau fauna. Ujung Kulon is one 
such area. This paper is an appraisal of the butterflies collected and 
seen there during the expedition. 


METHODS 


A total of 158 biologist-days was spent in the Ujung Kulon area from 
30 August-—23 September 1984. Slightly under half of these (73 biologist 
days) were on Pulau Peucang (Fig. 1), an island of ca. 440 ha separated 
from the peninsula by ca. 500 m of shallow water. The various sites 
visited on the W part of Ujung Kulon are shown in Fig. 1. They include 
a range of representative vegetation types: Pulau Peucang—pes-caprae 
formations, Ficus forest, and various edge habitats; Ujung Kulon—pes- 
caprae, mangrove, Radermachera forest, Arenga forest, forest edges, 
and Chrysopogon grassland. Plant nomenclature is based on papers in 
Flenley and Richards (1982). 

Most records are of butterflies collected by the authors, but many 
other expedition members also contributed specimens. TRN and MBB 
systematically searched the above-noted vegetation types to observe 
relative abundance of the species present. Examples of the taxa collected 
will be deposited in the Museum Zoologicum Bogoriensis, Bogor. Iden- 
tification was from published literature (including Corbet et al. 1978 
and papers referred to therein), and by comparison with collections in 
Bogor and the British Museum (Natural History), except for Hesperi- 
idae, which were identified by A. F. Atkins. Although Blower and van 
der Zon (1977) refer to “large entomological collections” from the area, 
we have not traced any previous lists of butterflies from Ujung Kulon. 


VOLUME 41, NUMBER | 31 


N 









PULAU 
PANAITAN 


Cre § 
UJUNG KUioN 
PENINSULA 2 ee 


JAVA 





Fic. 1. Western part of Ujung Kulon peninsula, Java, indicating areas (shaded) where 
collections were made and locations inentioned in text. Sunda Strait passes between 
Sumatra and Java, and the Krakatau Islands are in the center of the Strait 60 km N of 
Ujung Kulon. 


RESULTS AND DISCUSSION 


Although the butterfly fauna of Indonesia is reasonably well known, 
at least in gross terms, there are few lists of species in areas of particular 
biological significance. The fauna of the Krakatau Islands has received 
much recent attention (Yukawa 1984, Bush 1986, New et al. 1987), and 
Yukawa (1984) also noted 29 species from Pulau Panaitan, an island of 
slightly over 12,000 ha separated from Ujung Kulon by ca. 10 km of 
deep water, and not visited by us. The data presented here thus com- 
plement Yukawa’s account and, although the species list (Table 1) 


39 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Butterflies recorded in Ujung Kulon Peninsula and Pulau Peucang in 1984 
including records in Pulau Panaitan from Yukawa (1984) and incidence of species on the 
Krakatau Islands. 


Pulau ‘Pulau 
Ujung Peu- Panai- Kraka- 
Kulon cang tan taus 


Papilionidae 
Pachliopta aristolochiae antiphus (F.) 
P. a. adamas (Zinken) 
Atrophaneura coon coon (F.) 
Graphium agamemnon agamemnon L. 
G. doson evemonides Honrath 
G. macareus macareus (Godart) 
Papilio peranthus peranthus F. 
P. helenus engarius Doherty 
P. polytes? javanus Felder 
P. antiphates Cramer ssp.? 
P. memnon anceus Cramer 


xX 
xX 


wm MK PM 


PPM MO 
ars 


Pieridae 


Appias indra leptis (Felder) 

A. lyncida lyncida (Cramer) 

A. nero nero (F.) 

A. paulina (Cramer) 

Catopsilia pomona F. 

Ceporea idith (F.) 

Eurema blanda (Boisduval) 

E. hecabe (L.) 

Leptosia nina (F.) 

Gandaca harina harina (Horsfield) 
Pareronia valeria leona (Fruhstorfer) 
Saletara liberia panda (Godart) 
Hebemoia glaucippe (L.) 


ms 
mx rex 
PS KM PS PS PS OS 
rs 


PS PS PS PS PS PS PS PS OS OM 
rs 


Danaidae 


Euploea camerelzeman Butler 

. crameri Lucas 

. diocletianus F. 

. mulciber mulciber (Cramer) 

. m. donada Fruhstorfer 

. modesta Butler 

. leucostictos leucostictos Gmelin 
. tulliolus (F.) 

Radena juventa (Cramer) 
Tirumala limniace (Cramer) 


Baas 


es MM RM OK 
re 


Nymphalidae 
Agatasa franck Godart 
Athyma nefte subrata Moore f. neftina (Fruhstorfer) xX 
Cethosia penthesilea methypea (Butler) xX 
C. hypsea Doubleday 
C. sp.? xX 
Charaxes sp. 
Chersonesia rahria (Moore) 
Cirrhochroa tyche (C. & R. Felder) 
Cupha erymanthis lotis (Sulzer) 


x 
Cyrestis thermire Honrath 
pn a a 


va 


re rs 
rrr 
re 


VOLUME 41, NUMBER 1 33 


TABLE 1. Continued. 


Pulau — Pulau 
Ujung Peu- Panai- Kraka- 
Kulon cang tan taus 


Doleschallia bisaltide pratipa (C. & R. Felder) 
Euthalia evelina sikandi (Moore) 

E. mahadeva (Moore) 

Hypolimnas anomala anomala (Wallace) 
Lebathea martha malayana Fruhstorfer 
Lexias dirtea (F.) 

Neptis hylas (L.) 

N. sankara (Kollar) 

Pantoporia paraka (Butler) 

Phalanta alcippe (Stoll) 

Precis atlites L. 

P. almana javana C. Felder 

P. erigone (Cramer) 

P. hedonia ida (Cramer) 

P. iphita (Cramer) 

Tanaecia clathrata (Vollenhoeven) 

T. godartii (Gray) 

T. iapis (Godart) 

T. munda Fruhstorfer 

Terinos terpander ?teos (de Niceville) 


PA PS PK PS PS PM PS PS PS KS PS PS PS PPS PS 
rs 
ms 


Ps rs 


Satyridae 


Melanitis leda (L.) 

M. phedima (Cramer) 
Elymnias hypermnestra (L.) 
Lethe confusa Aurivillius 
Mycalesis horsfieldi (Moore) 
M. janardana Moore 
Ypthima horsfieldi Moore 
Faunis canens canens Hiibner xX 


PA PS PS PS MS 
axxo MM 


Lycaenidae 


Liphyra brassolis Westwood 
Arhopala antimuta C. & R. Felder 
Allotinus horsfieldii Moore 

A. subviolaceus C. & R. Felder 

A. unicolor C. & R. Felder 
Catochrysops panormus (C. Felder) 
C. strabo (F.) 

Chilades pandava (Horsfield) 
Drupadia ravindra Distant 
Eooxylides tharis distanti Riley 
Ionolyce helicon (C. Felder) 
Jamides bochus (Stoll) 

J. aratus (Stoll) 

J. celeno (Cramer) 

J. elpis (Godart) 

J. parasaturatus (Fruhstorfer) 

J. malaccanus (Rober) 

Lampides boeticus (L.) 

Miletus ?boisduvali Moore 

M. sp. X 


KX 


a a a a a oe a a 
va 
rr 


34 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Continued. 


_ Nacaduba pactolus (C. Felder) xX 
Neopithecops zalmora (Butler) 
Pithecops corvus Fruhstorfer 
Prosotas nora superdates Fruhstorfer 
P. dubiosa (Semper) 

Zizina otis (F.) 
Zizula hylax (F.) 


Hesperiidae 
Halpe pelethronix Fruhstorfer 
Koruthaialos rubecula namata Fruhstorfer 
Isma bononia bononia (Hewitson) 
I. obscura vulsina Evans 
Sancus (Psolos) fuligo fuligo (Mabille) 
Telicota colon vaja Corbet 
Zographetus (Ogygia) ogygioides Elwes & Edwards 
Salanoemia tavoyana Evans 
Acerbas anthea javanica Snellen 
Taractrocera? aliena aliena (Plotz) 
Potanthus confucius (C. & R. Felder) xX xX 


mx KM 


mx xX 


PK mM PS PX 


mr XK 


cannot be regarded as comprehensive, it will be of value for comparison 
with other sites in Indonesia as the fauna is progressively documented. 

In Table 1, the species are noted as occurring either on Ujung Kulon 
proper or on Pulau Peucang. Yukawa’s (1984) Pulau Panaitan records 
are also included, and incidence of species on the Krakatau Islands is 
indicated. 

At least 106 species of butterflies are here recorded from Ujung Kulon 
and nearby islands; the several noted merely as “‘sp.”’ are not included 
in this total because of possible overlap. This figure represents some 
18% of the 583 species recorded from Java (Yukawa 1984), and includes 
representatives of all families except Libytheidae (3 Javanese species) 
and Riodinidae. The relatively small size of Pulau Peucang rendered 
our coverage reasonably complete, and the 36 species recorded there 
are believed to well represent the butterflies then flying on the island. 

Vegetation of Ujung Kulon is diverse (Hoogerwerf 1970, Blower & 
van der Zon 1977, Hommel 1983), and offers a wide range of habitats 
for butterflies: beach forests with pes-caprae formations, mangrove 
swamp forests, freshwater swamp forests, and rain forest, and artificially 
maintained clearings and grasslands. 

Distribution of many butterfly species was limited, and clearly related 
to predominant vegetation types. The pes-caprae formations take their 
name from Ipomoea pes-caprae (L.) R. Br., a common creeper on 


VOLUME 41, NUMBER lI 35 


accreting tropical beaches growing in association with a mixture of 
grasses and herbaceous dicotyledons. The pes-caprae formations of both 
Pulau Peucang and Ujung Kulon were rich in Jamides spp., Cato- 
chrysops spp. and Eurema spp. Catochrysops strabo (F.) was the com- 
monest of the lycaenids in this association on both Pulau Peucang and 
Ujung Kulon, a finding which contrasts markedly with its apparent 
absence from Pulau Panaitan (Yukawa 1984) where its place appears 
to be filled by C. panormus (C. Felder). Both C. strabo and C. panormus 
were recorded on the pes-caprae formations on Krakatau, but both were 
relatively scarce there, and Jamides celeno (Cramer) and J. aratus 
(Stoll) were the predominant species. Further work is necessary to 
establish the seasonality of these species before a significant distribution 
pattern can be discerned. 

The narrow belt of mangroves at Cidaon supported few butterfly 
species. Those seen flying there, including Papilio helenus engarius 
Doherty and Melanitis phedima (Cramer), seldom settled, possibly 
having strayed from the nearby forest edge. 

The forests of Pulau Peucang and Ujung Kulon differ widely in 
character. Those of the former appear to be more mature, and are 
dominated by large individuals of Ficus pubinervis Bl. and strangler 
figs, which form a canopy at ca. 30-35 m. The canopy produced by 
these species is dense; little light reaches the forest floor, and there is 
very little undergrowth between the widely spaced trunks. Only Terinos 
terpander ?teos (de Niceville) and Melanitis leda (L.) were recorded 
within this forest. However, where trees had fallen, creating an opening 
in the canopy, Phalanta alcippe (Stoll) and Drupadia ravindra Distant 
were present. This forest is in some ways comparable to the Ficus 
pubinervis forest of Rakata, Krakatau. On Rakata the F. pubinervis 
forest is not so well established, and there is a dense undergrowth of 
pteridophytes and Smilax zeylanica L. The last is a forest creeper which 
has been suggested as the foodplant of the only deep-forest butterfly 
on Krakatau, Loxura atymnus fuconius (Stoll) (Bush 1986). Notably, 
neither L. atymnus nor S. zeylanica were recorded during our visit to 
Pulau Peucang. 

One similarity between the forests of Krakatau and Pulau Peucang 
is the lack of impact of footpaths. On the latter, the trunks of trees are 
so widely spaced that there has been no need to fell trees to create 
paths, and thus the canopy remains unbroken. Krakatau has no foot- 
paths; the only clearings are where trees have fallen. This contrasts with 
Ujung Kulon where the denser forest growth necessitated limited clear- 
ing for footpaths. The result is the creation of forest “rides”, where 
more light reaches the forest floor. If the butterflies found along these 
forest paths are used as an indication of the overall species composition 


36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


in forest areas, two points must be borne in mind. First, there is an 
increased butterfly population taking advantage of the flush of her- 
baceous growth along the footpath, and this might give a misleading 
impression of the density of butterfly populations. Second, there is the 
problem of dissociating those species which are true deep-forest but- 
terflies (and have been found by chance along the ride) from those 
species which are woodland-edge species and would not be found there 
but for the increased light availability. An impression of butterfly num- 
bers and species diversity in the true forest can only be obtained away 
from the paths. 

Butterflies which seemed to be characteristic of the Ujung Kulon 
forest interior were Pithecops corvus Fruhstorfer, Arhopala antimuta 
C. & R. Felder, Agatasa franck Godart (the last seen roosting high on 
tree trunks), and locally Lexias dirtea (F.) (found only on the higher 
slopes of Gunung Payung (480 m)). 

The forest edge habitat on Pulau Peucang was probably oversampled 
compared with other areas, as the clearing surrounding the park office 
bungalows where we were based was the nearest and easiest area to 
collect in. Pink and red-flowering garden shrubs attracted a variety of 
danaids including Euploea crameri Lucas, E. mulciber mulciber (Cra- 
mer) and Radena juventa (Cramer). Another common visitor to these 
shrubs was Papilio peranthus peranthus F., although it appeared to be 
scarce in the rest of the reserve, and was not recorded at all from Ujung 
Kulon. Graphium agamemnon L., Precis atlites L. and Neptis hylas 
(L.) were among other heliophilous species recorded in the clearing. 

Forest-edge areas on Ujung Kulon are a varied set of habitats deter- 
mined by the surrounding forest type. There is additional habitat di- 
versity created by stream banks. Many species were collected along 
forest paths, or in clearings where trees had fallen. Some of those that 
may be true forest species were Leptosia nina (F.), Allotinus spp., 
Lebathea martha malayana Fruhstorfer (?), and members of the Eu- 
thalia-Tanaecia complex. Species that seemed more light-demanding 
recorded along paths were Hebemoia glaucippe (L.), Appias spp., Atro- 
phaneura coon coon (F.) and Ypthima spp. Many Pieridae were cap- 
tured on flowers or at streamsides; the most abundant were Eurema 
spp. and Catopsilia pomona F., the latter migrating at the time of our 
visit (New et al. 1985). The various species of Appias were all taken in 
open glades or at the edge of the forest where, like H. glaucippe, they 
visited flowering shrubs. Although some of the pierids were caught near 
streams, there were no marked concentrations of butterflies on stream 
banks. 

In Ujung Kulon the extensive Chrysopogon grassland communities 
maintained for herds of banteng, Bos javanicus (d’ Alton), did not ap- 


VOLUME 41, NUMBER 1 37 


TABLE 2. Numbers of butterfly species shared between areas of western Java and the 
Krakatau Islands. 


Ujung Kulon Pulau Peucang Pulau Panaitan Krakatau Islands 
Ujung Kulon — 24 iM 22 
P. Peucang — — 6) 15 
P. Panaitan — — — 9 
Total 85 36 29 54 


pear to support a rich butterfly fauna. This, however, may have been 
due to cloudy conditions when collecting was carried out on the main 
grassland area at Cidaon. The abandoned field systems around Cibunar 
were rich in flowering herbs such as Eupatorium odoratum L., which 
attracted Neptis spp. and Pachliopta aristolochiae (F.). 

The eight species of Hesperiidae recorded from Ujung Kulon were 
captured along forest tracks or at the forest margin. None was taken 
on the grassland expanses of Cidaon or Cibunar. These are all woodland 
or semi-woodland species. The eight species of Hesperiidae from Krak- 
atau were caught in grassland or beach habitats, and habitat preferences 
may thus account for the lack of overlap of species between the two 
localities. 

One butterfly recorded regularly on Krakatau (and also at Carita, 
W. Java) was Troides helena (L.). However, neither T. helena nor the 
closely related T. cuneifer (Oberthiir) were recorded from Ujung Kulon 
or associated islands. Delias spp., which might have been expected in 
the forests at Ujung Kulon, were notably absent. This genus was also 
absent from Krakatau, but there the reason is likely to be the absence 
of the foodplants (Loranthaceae). 

Degrees of overlap of species found on Ujung Kulon, Pulau Peucang, 
Pulau Panaitan, and the Krakatau Islands are shown in Table 2. Many 
of the 54 species found in recent collections from Krakatau (New et al. 
1987) were not found in the Ujung Kulon area. 

Tables 1 and 2 should be interpreted with caution as the survey of 
Ujung Kulon was far from complete, and the low values obtained from 
Pulau Panaitan suggests that it, too, may have been undersampled. It 
is likely that all the species found on Pulau Peucang and Pulau Panaitan 
are present on Ujung Kulon even though they were not recorded during 
this census. From the data available, the degrees of species overlap of 
Ujung Kulon with Pulau Peucang and Krakatau are almost equal. How- 
ever, the species are not the same in each case. Fourteen species have 
been recorded as common to all three localities. These species are either 
highly mobile-migratory or inhabit the coastal vegetation and pes- 
caprae associations. Radena juventa, Neptis hylas and Catopsilia po- 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


mona which are geographically widespread or migratory, tend to be 
polyphagous, and are, therefore, likely to be successful colonizers. Species 
that may be more specific in their foodplant requirements, including 
the species of the pes-caprae formations, must rely on their habitat 
being present at a new site if colonization is to be successful. In coastal 
species, two factors aid dispersal to islands. First, they are likely to be 
blown out to sea, and second, the plants adapted to beach environments 
are likely to be among the first colonizers of an island where plant 
recolonization is taking place. Thus the foodplants are available for the 
coastal butterfly species on their arrival. Woodland butterfly species 
may have to arrive many times before they find a suitable habitat. This 
might help to explain the apparent paucity of woodland butterfly fauna 
on the offshore islands considered in this paper compared to the diverse 
species list for forest edges and coastal environments. 

Particular families also overlap. As already noted, there was no over- 
lap between the Hesperiidae of the Krakatau Islands and those of Ujung 
Kulon, and yet five out of eight species of satyrids found on Ujung 
Kulon were also found on Krakatau. In all the areas visited the com- 
monest member of the Satyridae was Melanitis leda. 

Many Lycaenidae were captured in small numbers, and because of 
the existence of species complexes, exact identification is not always 
clear. A single specimen of Liphyra brassolis Westwood is of interest, 
as this species is apparently rare in much of Indo-Malaya (Corbet et 
al. 1978). Miletinae were represented by species of Allotinus and Mi- 
letus, and Polyommatinae were by far the most diverse group of ly- 
caenids present. The species shared with the Krakatau Islands are pre- 
dominantly those associated with lowland Leguminosae, including the 
pes-caprae coastal vegetation which is also an important butterfly hab- 
itat on the Krakatau Islands (Bush 1986, New et al. 1987). Theclinae 
were relatively scarce, although one species of the complex genus Ar- 
hopala was found; the single species of Eooxylides and Drupadia are 
both widely distributed in Sundaland, but have not been found on 
Krakatau. 

Hesperiidae can be identified only with some reservations. Only 
females of Taractrocera a. ?aliena (Plotz) were taken, for example, and 
male genitalia are necessary for confirmation of species identification. 
Most of the species records are based on single individuals. 


CONCLUSIONS 


All the butterfly species captured are likely to be resident in the area, 
and the vegetational diversity is clearly sufficient to support an enor- 
mous spectrum of Lepidoptera. The range from mature forest to cleared 
ground provides a diversity of successional stages. The present conser- 


VOLUME 41, NUMBER 1 39 


vation policy of maintaining cleared areas for grazing by banteng helps 
to foster such diversity. It has been suggested that there should be a 
management policy to create forest glades to provide improved grazing 
for the Javan rhinoceros. Such areas would also help to maintain but- 
terfly diversity, but felling would have to be done with sensitivity to 
leave areas of mature forest for species that are dependent on deep- 
forest habitat. However, as Hommel (1983) suggests, clearings are pro- 
duced naturally when giant forest trees fall. This natural process, com- 
bined with maintenance of the footpath system, could provide habitats 
similar to those advocated by Schenkel et al. (1978) without recourse 
to more active management. Further study is needed to determine the 
distribution of particular butterfly species on Ujung Kulon, but this 
brief survey, although restricted to the western region of the park, 
indicates that some species may be both rare and localized. Butterfly 
diversity in this area may well depend on the maintenance of the 
greatest possible range of vegetation types. 


ACKNOWLEDGMENTS 


We thank I. W. B. Thornton (La Trobe University) for leading the joint La Trobe 
University-Bogor Zoological Museum Expedition to the Krakataus and for reading a draft 
of this paper; S. Adisoemato (Museum Zoologicum Bogoriense) for logistic support; and 
our colleagues who collected some of the specimens. We also acknowledge permission to 
work in the Ujung Kulon National Park granted by the Pelindungen dan Pengawetan 
Alam, and thank Lembaga Ilmu Pengetahuan Indonesia for allowing us to undertake 
scientific work in Indonesia. 


LITERATURE CITED 


BLOWER, J. H. & A. P. M. VAN DER ZON. 1977. Proposed Ujung Kulon National Park 
including Gunung Honje, Pulau Peucang and Pulau Panaitan. Management Plan 
1977-1981. Field Report of U.N.D.P./F.A.O. Nature Conservation and Wildlife 
Management Project. INS/73/013. FAO, Rome. 90 pp. 

BusH, M. B. 1986. The butterflies of Krakatoa. Entomol. Monthl. Mag. 122:51-58. 

CoRBET, A. S., H. M. PENDLEBURY & J. N. ELIOT. 1978. The butterflies of the Malay 
Peninsula. Kuala Lumpur, Malayan Nature Society. 578 pp. 

FLENLEY, J. R. & K. RICHARDS. 1982. The Krakatoa Centenary Expedition. Final Report. 
University of Hull, Dept. of Geography, Misc. Ser. 25. 196 pp. 

HoMMEL, P. W. F. M. 1983. Ujung Kulon vegetation survey—Preliminary results 
including a landscape-ecological map. World Wildlife Fund. 84 pp. 

HOOGERWERF, A. 1970. Udjung Kulon. The land of the last Javan rhinoceros. E. J. Brill, 
Leiden. 512 pp. 

NEw, T. R., M. B. Bus, I. W. B. THORNTON & H. K. SUDARMAN. 1987. The butterfly 
fauna of the Krakatau Islands after a century of colonisation. Phil. Trans. Roy. Soc. 
Lond., B (submitted). 

NEw, T. R., G. S. FARRELL, N. W. Hives & P. A. HORNE. 1985. An early season 
migration of Catopsilia pomona (Lepidoptera: Pieridae) in Java, Indonesia. J. Res. 
Lepid. 24:84-85. 

SCHENKEL, R., L. SCHENKEL-HULLIGER & W. S. RAMANO. 1978. Area management for 
the Javan rhinoceros (Rhinoceros sondaicus Desm.) A pilot study. Malayan Nature 
J. 31:253-275. 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


THORNTON, I. W. B. (ed.). 1985. 1984 zoological expedition to the Krakataus. Prelim- 
inary Report. La Trobe University, Department of Zoology. 57 pp. 

THORNTON, I. W. B. & N. J. ROSENGREN. 1987. Zoological expeditions to the Krakataus, 
1984, 1985: General introduction. Phil. Trans. Roy. Soc., B (submitted). 

YUKAWA, J. 1984. Geographical ecology of the butterfly fauna of the Krakatau Islands, 
Indonesia. Tyo to Ga 35:47-74. 


Received for publication 22 July 1986; accepted 6 November 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 41-44 


THE IDENTITY OF CYCLOPIDES PAOLA PLOTZ 
(HESPERIIDAE) 


Davip L. HANCOCK 


5 Northampton Crescent, Hillcrest, Bulawayo, Zimbabwe 


ABSTRACT. Cyclopides paola Plétz, described from Angola, is placed in the com- 
bination Kedestes nerva paola (Plotz), new status. Kedestes protensa Butler, previously 
regarded as a subspecies or synonym of K. paola, is reinstated as a species. Records of K. 
nerva (Fabricius) from Zimbabwe and Kenya are rejected. 


Additional key words: Africa, taxonomy, Kedestes protensa, K. nerva. 


Ever since Pl6tz (1884) described Cyclopides paola from Angola, its 
identity has been a source of confusion. Its generic placement was 
questioned by Holland (1896), while Swinhoe (1908) referred it to 
Kedestes Watson, and published a color figure based on Plétz’s original 
unpublished illustration, noting that it could belong to either K. nerva 
(Fabricius) (=tucusa (Trimen)) or K. protensa Butler. Evans (1937) 
regarded it as conspecific with K. protensa, with the latter a subspecies. 
The presence of a complete marginal line on the underside of the 
hindwing in the figure of paola, plus Evans’ association of the taxon 
with K. protensa, led Hancock and Gardiner (1982) to synonymize the 
two, since Evans’ (1937) interpretation of paola as a subspecies was at 
variance with available material. As used by Pennington (1978), the 
name paola is now known to refer to three separate species: K. protensa, 
K. michaeli Gardiner & Hancock and K. monostichus Hancock & 
Gardiner (Hancock & Gardiner 1982). 

The capture in NW Zambia of a fresh pair of K. nerva (Ikelenge, 5 
May 1983, D. Heath; in A. Heath Collection, Cape Town), and a re- 
examination of the male from Mwinilunga recorded and illustrated in 
Hancock and Gardiner (1982) (in Natural History Museum of Zim- 
babwe, Bulawayo), revealed minor differences in wing pattern between 
them and typical examples of nerva from South Africa, although there 
are no discernible differences in male genitalia. These Zambian males 
of K. nerva agree more closely with the figure of paola given by Swinhoe 
(1908) than do specimens of K. protensa from the same area (figured 
by Hancock & Gardiner 1982 as paola), particularly with regard to the 
elongate white spot in space 2, the pale area beyond the cell on the 
forewing underside, and the ground color and distribution of the black 
spots on the hindwing underside. In size too, the Zambian nerva agree 
with the type of paola, while protensa is a distinctly larger species. The 
marginal line on the hindwing underside is a series of streaks, inter- 
rupted at the veins, and not a continuous line as shown in Swinhoe’s 
(1908) figure. 

The original description of paola by Plétz (1884) is sketchy, and the 


42 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


marginal line on the hindwing is not mentioned. The type is apparently 
lost, and subsequent interpretations of the species appear to be based 
on Plétz’s unpublished plate (the present whereabouts of which, if still 
extant, is unknown to me), and on the reproductions by Swinhoe (1908) 
and Aurivillius (1925). The continuous marginal line on the hindwing, 
evident in figures of the latter two authors, is apparently an error. It 
may well be present also on the original figure, as the line appears 
continuous unless examined closely; the streaks are longer and not as 
distinct as in typical examples of K. nerva. 

Kedestes protensa (=K. paola of Hancock & Gardiner 1982) and K. 
nerva differ in wing size and in shape of male genitalia, particularly 
the serrate apical area of the harpe, which is narrower and projecting 
in K. nerva. In pattern, apart from the hindwing marginal line, the 
two species differ in the arrangement of the black spots in the postdiscal 
band on the hindwing underside; the inner row of spots is distinctly 
W-shaped in K. nerva, V-shaped in K. protensa. The taxon paola differs 
from both these species in the reduced pale spot in space 1b of the 
forewing, but it agrees with K. nerva in male genitalia and other pattern 
characters. The ground color of the hindwing underside is browner in 
paola than in the other species, but this color is approached in some 
examples of nerva from South Africa, although an orange tint is usually 
evident. Accordingly, K. paola is placed here as a subspecies of K. 
nerva. Illustrations of these taxa may be found in Hancock and Gardiner 
(1982) and Pennington (1978), but in both cases Zambian examples of 
K. protensa are misidentified as K. paola. Hancock and Gardiner (1982) 
also figured male genitalia. 

The establishment of the true identity of K. paola necessitates an 
adjustment to the nomenclature of K. nerva and K. protensa from that 
given in Hancock and Gardiner (1982). Kedestes paola is separable as 
a subspecies of K. nerva on pattern characters. The revised nomencla- 
ture is presented below. 


Kedestes nerva nerva (Fabricius) 


Hesperia nerva Fabricius, 1793:340. Type from “Indiis”, locality error, recte Natal, South 
Africa. 
Pyrgus tucusa Trimen, 1883:359. Type 2 from Natal, South Africa. 


This subspecies appears to be restricted to South Africa, being re- 
corded from the provinces of Natal and Transvaal. Records of Zambia 
(Hancock & Gardiner 1982) belong to the following subspecies, while 
those from Zimbabwe (Pinhey 1949, Pennington 1978) and possibly 
Kenya (Evans 1946) appear to belong to K. michaeli. Examination of 
a female of K. michaeli from Zimbabwe (Bromley, 23 August 1967, E. 
O. Martyn; in Natural History Museum of Zimbabwe, Bulawayo), iden- 


VOLUME 41, NUMBER 1 43 


tified by the collector as K. nerva, illustrates the confusion that existed 
between these two species, making the untraced Salisbury record of K. 
nerva noted by Hancock and Gardiner (1982) very unlikely. Evans 
(1946) noted that the Kenyan specimens he examined were not quite 
typical of nerva, and these and the Zimbabwe records are therefore 
rejected. 


Kedestes nerva paola (Plotz), new status 


Cyclopides paola Plotz, 1884:392. Type 6 from Angola. 


This subspecies is known from Angola and the Mwinilunga district 
of NW Zambia. It differs from typical nerva in the reduced white spot 
in space 1b on the forewing, and in the browner ground color and 
better developed marginal streaks on the hindwing underside. Evans 
(1937) appears to have confused this taxon with K. michaeli, and all 
his records from Zambia, Zaire, and Angola probably belong to this 
latter species, suggested by the presence of only a single cell spot on 
the forewing. This cell spot is divided into two in nerva and protensa. 


Kedestes protensa Butler, revised status 


Kedestes protensa Butler, 1901:59. Type ¢ from N. Nigeria. 

Kedestes chacoides Gaede, 1915:126. Type 6 from Busantare, Cameroon. 

Kedestes paola protensa Butler; Evans, 1937:84. 

Kedestes paola; Pennington, 1978:fig. 724 ii; Hancock & Gardiner, 1982:120. Misiden- 
tifications. 


This species is known from Sierra Leone, Nigeria, Cameroon, Ugan- 
da, S Sudan, and NW Zambia. It appears to prefer swampy areas of 
grassland, a moister habitat than that apparently preferred by K. nerva. 


LITERATURE CITED 


AURIVILLIUS, C. 1925. Hesperiidae. In Seitz, A. (ed.), The Macrolepidoptera of the 
world. Vol. 13. Kernan, Stuttgart. 

BUTLER, A. G. 1901. On a collection of butterflies made by George Migeod, Esq., in 
northern Nigeria between September 1899 and January 1900. Ann. Mag. Nat. Hist. 
(7)8:57-60. 

Evans, W. H. 1937. A catalogue of the African Hesperiidae. British Museum (Natural 
History), London. 

1946. Revisional notes on African Hesperiidae. Ann. Mag. Nat. Hist. (11)13: 
641-648. 

FABRICIUS, J. C. 1793. Entomologica systematica. Vol. III. 

GAEDE, M. 1915. Neue Afrikanische Lepidopteren des Berliner Zoologisches Museums. 
Int. Entomol. Z. 9:105, 106, 109-112, 125-126. 

Hancock, D. L. & A. J. GARDINER. 1982. The Kedestes nerva group of species (Lep- 
idoptera: Hesperiidae). Arnoldia Zimbabwe 9:105-123. 

HOLLAND, W. J. 1896. A preliminary review and synonymic catalogue of the Hesper- 
iidae of Africa and the adjacent islands. Proc. Zool. Soc. Lond. 1896:2—107. 

PENNINGTON, K. M. 1978. Pennington’s butterflies of Southern Africa. Ed. C. G. C. 
Dickson with D. M. Kroon. Donker, Johannesburg. 670 pp. 





44 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


PINHEY, E. C. G. 1949. Records of southern Rhodesian butterflies (Rhopalocera). Occ. 
Pap. Natl. Mus. Sthn. Rhod. 2:276-341. 

PLOTz, C. 1884. Die Gattung Cyclopides Hibn., und ihre Arten. Stett. Entomol. Z. 45: 
389-397. 

SWINHOE, C. 1908. On the species of Hesperiidae, from the Indo-Malayan and African 
regions, described by Herr Plétz, with descriptions of some new species. Trans. 
Entomol. Soc. Lond. 1908:1-36. 

TRIMEN, R. 1883. Descriptions of twelve new species of South African Lepidoptera 
Rhopalocera. Trans. Entomol. Soc. Lond. 1883:347-363. 


Received for publication 7 July 1986; accepted 10 November 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 45-64 


MATE LOCATION BEHAVIOR OF THE LARGE SKIPPER 
BUTTERFLY OCHLODES VENATA: FLEXIBLE 
STRATEGIES AND SPATIAL COMPONENTS 


R. L. H. DENNIS 
The Manchester Grammar School, Manchester M13 OXT, U.K. 


AND 


W. R. WILLIAMS 
Computer Centre, Science Laboratories, Durham DH1 3LE, U.K. 


ABSTRACT. Male Ochlodes venata were studied in lightly wooded heathland at 
Lindow, Cheshire, U.K., using a combination of transects and more detailed observations 
of movements and behavior in a woodland clearing. Males locate mates by perching and 
patrolling but are also opportunists and approach females when feeding. Behavior is 
flexible, and varies with time of day, weather, season, and location. On clear, sunny days, 
males show a significant bias for patrolling during mid- and late morning, and for perching 
during early morning and afternoon. Switches then from one activity to another are not 
entirely related to temperature and energy levels, and may depend on availability of 
females, males perching when females are relatively scarce. Scent seems to play a prom- 
inent part in the location of females during patrolling. Defended territories are located 
at topographic vantage points (at habitat edges and path junctions), and have a biased 
distribution to areas where female resources (nectar, oviposition sites) are also available. 
Territories and even the same perches (specific locations where the insects alight) within 
them are used by different individuals in the same and different years. 

Additional key words: Hesperiidae, England, territoriality. 


In the process of obtaining mates, male butterflies are described by 
Scott (1974) as adopting one of three strategies: perching or waiting 
for them, patrolling or actively seeking them in flight, and locating 
them with the use of long-distance pheromones. The last is regarded 
as of rare occurrence and understood to be a proximate cue (Scott 1972), 
vision dominating long-distance communication. But Heliconius erato 
males locate preemergent females using chemicals emitted by the pupae 
(Bellinger 1954). Perching males select characteristic sites, distinct in 
surface, height, and situation where they bask in the sun and launch 
themselves at passing insects (Baker 1972, Davies 1978, Dennis 1982a, 
Bitzer & Shaw 1979, 1983). Patrolling males typically cover much larger 
areas but differ in the extent and nature of the areas covered (Courtney 
1980, Dennis 1982b, Peachey 1980). Scott (1974) regards these processes 
as distinct; when perching occurs the females necessarily find males, 
but patrolling males actively find females. Perching has been likened 
to territoriality by Baker (1972), since defense of some “resource” is 
involved, but this is denied by Scott (1974). 

Recently, mate location behavior has been shown to be more varied. 
Perching and patrolling provide alternative strategies in some species 
depending on demographic trends (distribution of females, male-female 


46 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ratio) and environmental conditions (sunshine and temperatures, time 
of day, habitat topography, physical resources) (Scott 1975, 1983, Dennis 
1982a, Shreeve 1984, Rutowski 1983, Wickman 1985a). Moreover, it 
seems that many butterflies adopt intermediate strategies, involving 
intermittent flight, basking, and feeding bound only to loose areas and 
without territories, where males are constantly vigilant for females 
(Dennis 1982a, Morton 1985). 

The various aspects of mate location in butterflies are far from re- 
solved. The present paper on Ochlodes venata (Bremer & Grey) re- 
sponds to the need for comparative data. In particular, it addresses 
issues of territoriality, switches in mate location activity, and siting and 
structure of territories. O. venata is single-brooded in Britain and flies 
from the latter half of June into late July (Heath et al. 1984). 


METHODS 


The study was conducted intermittently over a three-year period 
(1983 to 1985) at Lindow Common near Wilmslow in Cheshire, U.K. 
(grid reference SJ834812), a 16-ha site of special scientific interest, 
comprising patches of dry heath (Calluna vulgaris with Ulex spp., 
Vaccinium myrtillus, and Deschampsia flexuosa over podsol soils on 
fluvioglacial sands) and wet heath (Calluna and Erica tetralix on 
permanently wet peaty soils over boulder clay), encircled by Betula- 
Quercus woodland. The site is intensively used for recreation and is 
crisscrossed by paths and tracks. 

Data were obtained by recording behavior and location of adults 
along repeated transects over fixed routes (Pollard 1977, 1979). Tran- 
sects were established at two scales, one covering the variety of vege- 
tation zones on the common, the second limited to a small clearing. 
The clearing was divided down the center into two zones, and obser- 
vations made from the wooded margin. Transects were walked at a 
standard pace 6 times over 20 min in each hour. The clearing comprises 
an open space some 25 m by 18 m divided into 2 zones of Molinia 
caerulea and Calluna, and surrounded by birch canopy and furze. 
Spatial data were plotted on an Ordnance Survey 1:2,500 base and on 
a detailed plan of the clearing (Fig. 1). More intensive observations 
were made using a cassette recorder relating behavior and movements 
of adults to coded landmarks. Individuals were followed as long as 
possible (minimum of 3 min), and activities divided into the following 
categories: resting, basking, feeding, flight, interspecific interaction, 
conspecific interaction, attempted courtship, and inspection. To these 
were added locational data and as much interpretation as feasible. 
Capture of individuals for marking was found to affect their behavior, 
and for these observations each male required a unique mark. The 


VOLUME 41, NUMBER 1 47 


Th Zeke 1516 17 18 
25 


24 

23 

22 

eg 21 

20 

19 

eer 18 
Gee 

Seas 17 

16 


© Betula 


Quercus 
0-5 HEIGHT (m) 


Molinia caerulea 
Ulex europaeus 
Erica tetralix 


Calluna vulgaris 


0 
1% 95) 16 47 46, 





Fic. 1. Topography, vegetation, and shading of the clearing used for observations on 
territoriality. The scale around the edge of the map is also used as a coordinate system 
for Figs. 5, 6, 7 and 8. 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Counts of male O. venata for different habitats and associated flora from 
9 transects on Lindow Common. 


F % 1 % T 
Habitat 
Woodland cover 0 0 26 
Woodland glades & fringe 132 68 42 
Open sward 43 a, 27 
Open track 20 10 5 
Woodland bramble 0 0 5) 
Associated flora for open areas 
Calluna i 4 30 
Molinia 156 96 70 
Tall-herb grassland 11 5 9 
Bramble 97 50 13 
Dry heath 10 5 19 
Wet heath 11 6 9 


F, number of adults; % I, percent of insects recorded on transects; % T, percent of transect associated with the habitat 
and flora. Based on 195 insects. Length of transect 2.24 km. 


marking of individuals varied sufficiently for them to be traced follow- 
ing interactions with other males. Observations on particular individuals 
ceased if there was doubt as to their identity. All cassette recordings 
and transect observations were carried out in cloud-free conditions with 
shade temperatures above 20°C. As part of the program to determine 
the factors in the selection of perch sites, an artificial landmark (a white 
plastic bag) was used in the clearing, alternated with the natural back- 
ground for identical observation periods. 


RESULTS 
Distribution of Adults on Lindow Common 


Males have a clustered distribution on the common. They are not 
found under woodland shade (Table 1, x2, = 69.1, P < 0.001) and, in 
open areas, show a bias for wet heath, Molinia ea are. and tall-herb 
grassland as opposed to Calluna and dry heath (Table 1, x2.) = 35.5, 
P < 0.001). There is no preference for clearings and open spaces sur- 
rounded by woodland compared to open areas extending beyond wood- 
land (x?q)= 1.78, P > 0.1). Edge sites are preferred, along tracks in the 
woodland or out on the open heath, at path junctions, in woodland 
clearings, and along the woodland fringe, males congregating at bound- 
aries between tall and short vegetation rather than in open expanses of 
low growth (comparison with bramble areas _ excluded, 
X°u)= 35.1, P < 0.001). However, males significantly “clump” on bram- 
ble bushes (Rubus spp.) most of which are also located by the side of 
tracks or at path junctions (x24) = 223, P < 0.0001), but ignore bramble 
under dense woodland cover be = 47.7, P < 0.001). 


VOLUME 41, NUMBER 1 49 






ES Factors evenly mixed 
All = 20% & = 40% 
= One factor very smal! 

20% 


One factor dominant 
— 60 % 


WAVAVAVACS 
DPSS INIVIN) 0 


0 
100 Feed 0 


% 


Fic. 2. Behavior of 54 male O. venata observed for at least 3 min each (mean time = 
405 sec, standard deviation 249 sec) recorded in triangular co-ordinates for basking, flight 
and feeding. Percentage interactions are overlaid as proportional symbols. Inset diagrams: 
I, inactive; M, mobile; F, feeding. 


The female distribution is very similar to that of the males. On 
Lindow, O. venata larvae feed mostly on Molinia; females regularly 
oviposit on this grass and have been reared on it (Bink 1985). 


Modes of Behavior 


Modes or patterns of behavior are made up of individual acts of 
behavior. This is evident from placing 54 males in triangular coordinates 
for flight, basking-resting, and feeding. Several distinctive features 
emerge (Fig. 2). With three exceptions, males are restricted to areas 
on the graph where one activity is virtually dominant. Males do not 
behave randomly but bask, fly, or feed for extended periods, although 
grades of activity occur between flight and feeding and between basking 
and flight (far less between feeding and basking). Conspecific inter- 
actions (skirmishing between males and attempted courtship) occur for 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


each of these activities, which suggests that the butterfly is opportunistic 
in mate location. However, interactions are more frequent when males 
are basking. We argue that three distinctive types of behavior exist in 
male O. venata: perching, patrolling, and feeding. These behaviors are 
perhaps better visualized in the form of real-time plots of activity (Fig. 
3), and become easy to distinguish using the spatial components of each 
behavior pattern, as will become clear below. 

The perch behavior (Fig. 3a) is restricted to small areas, where a 
vantage point is adopted. Here, male O. venata bask in the sun and 
“sortie” after passing objects. Response to the latter varies dramatically 
from rapid inspections to violent spirals and chases depending on (i) 
the insect involved, (ii) the sex of the intruder if conspecific and (iii) 
prior ownership of the area. Conspecifics induce much greater activity 
than nonconspecifics, and contested ownership of the space leads to the 
most violent aerial activity, during which the impact of the males can 
be heard. Interactions between male O. venata are significantly longer 
when one of the males is territorial than when both are feeding or 
patrolling (¢ = 3.98, P < 0.001) (Table 2) (compare Wickman & Wik- 
lund 19838, Wickman 1985b). Passing females trigger attempted court- 
ship, though males are usually easily put off by fecund females, which 
flutter their wings and, having landed, on occasion, raise their abdo- 
mens. We suggest that some chemical deterrent is released. Males re- 
peatedly return to perches in the same small area, which distinguishes 
this activity from patrolling. Perches are typically low vegetation, small 
seedlings of oak or birch, bramble leaves, occasionally herbs or grass 
stems, which barely overtop the ground vegetation. Only rarely are 
perches more than 1 m above the ground adopted, but the butterfly 
will rest (become inactive) on vegetation above this level. Males feed 
least during perching; apart from interactions, flight is restricted to 
short investigative patrols (as distinct from continuous patrolling) within 
and immediately beyond the area occupied. 

At perch sites, extended interactions occur that are not simply the 
result of the resident male investigating the sex of the intruder and the 
intruder attempting to escape from a would-be predator (Scott 1974). 
Not only is female response to male approaches very different (many 
land and engage in mate-refusal posture), but this would not explain 
why intruding males (recognizable from wing marks) return after what 
can only be described as extremely violent aerial combat, and why such 
sorties take place back and forth over the area involved. The relation 
between the aggression of the incumbent and the inclination of the 
intruder to leave attests further the defensive role of interactions. Spe- 
cific tolerance thresholds need to be exceeded before intruders leave 
an area, and similarly determine whether incumbents stay or abandon 


VOLUME 41, NUMBER 1 51 


MALE —9: 9-7- 83, 11°47 











YM MM 








rie Resting - Basking Skirmishing 0 30 
—— 4 Liuiitiiitiiis Secs 
= Flight V4. Mttempted courtship 

mum =Feeding 


Fic. 8. Real-time plots of changing behavior in three male O. venata engaged in (a) 
territorial activity, #9, 1147 h, 9 July 88, (b) patrolling, #40, 1009 h, 4 July 85, (c) feeding, 
#8, 1187 h, 9 July 83. 


territories. To some extent it was possible to simulate these responses 
using a net. Perching males that are merely disturbed by passing a 
closed net over them invariably return to the perch; those caught in 
the net bag and released in one gentle move usually return or retire to 
an area nearby, and return within 10 seconds. Males caught in a sweep 


52 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


- TABLE 2. Duration of interactions between male O. venata and other insects. 





Interaction Statistics* Perching Patrolling and feeding 

Between males X em 4.3 

SE 0.7 0.4 

n 73 28 
With other insects? X 2.4 2.5 

SE 0.1 0.3 

n 76 16 
Attempted courtship x 3.8 3.8 

SE 0.7 0.3 

n 1 43 


4X, mean time (seconds); SE, standard error; n, sample size. 
> Other insects include Maniola jurtina, bees, and flies (mostly syrphids). 


of the net and then released exit in a straight line at high speed never 
to return (Fisher exact test, P < 0.001) (Table 3). This not only has 
connotations for territoriality but also for mark, release, and recapture 
work with O. venata. Finally, O. venata males reveal distinct signs of 
pugnacity to other insects, such as syrphids, which hover in the area, 
effecting direct onslaughts rather than gentle investigative flights of 
which they are capable. ; 

Patrolling (Fig. 3b) is distinguished by extended and spatially un- 
bounded flights. The flight can be very slow, at any height up to 3 m, 
but is usually very low, where the butterfly engages in weaving, scan- 
ning, hovering, often circular inspections interspersed with infrequent 
halts (physical inspections), but always remaining close to vegetation, 
skimming over its surface. Flight can also be much faster, several 
m/second, apparently when the butterfly is changing location, and cues 
are weak. Patrolling behavior is intensive and seems to be influenced 
strongly by scent. One male was observed searching a 1 m tall birch 
seedling systematically for 15 min, weaving in and out of the twigs and 
leaves, before it found the female. The female left the clearing followed 
by the male. On other occasions, persistent searches by males of small 
areas (<0.5 m?) in circuits were witnessed where females were found 
eclosing. 

Males feed while patrolling, but for short sessions. They also bask 
and rest, especially after extensive flight, although platforms differ from 
territorial perches (occurring at a greater range of heights up to 3 m; 
territorial perches are below 1 m), and are used once only. Skirmishes 
with conspecifics take place; some of these may well be short inspections, 
but many are sharp interactions which effect spacing of individuals. 

Feeding behavior (Fig. 3c) is equally distinctive, involving short 
feeding sessions interspersed by “‘hops’” or short flights from flower to 
flower. Short periods of basking and resting occur as do longer flights 


VOLUME 41, NUMBER 1 53 


TABLE 3. Effect of artificial disturbance on territorial male O. venata. 


Disturbance level Direct return Delayed return _ Rapid, linear exit 
Closed net swept above insect 10 0 0 
Gentle capture 6 8 2 
Determined capture 0 6 7 


as the butterfly relocates to new resources. Mate location is opportunistic 
rather than directly sought, non-conspecifics being ignored, and skir- 
mishes with conspecifics being brief but pugnacious or investigative. 
Attempted courtships also occur frequently, males tending to harrass 
nonreceptive females feeding on the same resource. The length of each 
feeding episode depends much on the nectar source, and is longer on 
Rubus spp. and Hierarchium than on Erica tetralix. 

The above three modes of behavior are distinctive and facilitate the 
classification of the 54 individuals in the ternary graph. Even so, only 
occasionally was it possible to follow males long enough to note switches 
from one behavior to another, although a number of males were seen 
to abandon territories, usually after prolonged periods of inactivity 
when intruders were lacking. 


Switches in Mate Location Activity 


Typically, males patrolled in the morning and perched in the after- 
noon (transect data over heath: x?,) = 5.8, P < 0.02; observations in 
clearings: x2, = 33.8, P < 0.001). The number of males engaging in 
territoriality increased in the afternoon in areas lacking nectar (x?) = 
13, P < 0.001) and in areas having it (x?) = 4.8, P < 0.05). This pattern 
was influenced by weather. During cloud-free days (Fig. 4), males perch 
early but switch to patrolling after 1000 h. In the afternoon, there is a 
reversion to perching, and some new territories are established. How- 
ever, in favorable locations, territorial males can be found throughout 
the day, as in the clearing (Fig. 1). Similarly, patrolling males can be 
found in the afternoon, but there is typically a great reduction in 
patrollers in the wet heath areas during afternoon. This pattern was 
much affected by weather; after overcast mornings, males patrolled in 
the afternoons. The marked diurnal pattern may also break down during 
the season. Preliminary observations point to a bias towards perching 
at the outset of the flight season and at the very end when unmated 
females are scarce. On 23 July, an equal number of perching males 
were recorded in the morning and afternoon. Other factors can lead 
to a breakdown of the typical model. During 1985, emergence was 
delayed by cloudy and cool weather during the last week of June. When 
conditions changed on 1 July, an abundance of fresh males and females 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


54 
% Time Number of Interactions 
100 ass 20 

/ wie a 18 

80 { a cal 
i 14 

60 i II 12 

i 10 

40 f 8 

i ‘ 

OM I 4 

\ 2 


176 19:7 22-0 233 24:4. 25:2 25:4) | 267g Zeee 


Numbers in Activity 












































24 
22 | a 
20 
te Perchers 
16 | | (im PATROLLERS fi \ 
14 me 
12 lh i > 
10 == —— 
5 a 
6 
4 
2 





9 (LO geet ia ake es 122 eatho) 2 3 4 5 (h) 
Time of Day 


Fic. 4. Diurnal changes in male O. venata behavior in the clearing for clear sky 
conditions on 7 July 1984. Top: Percentages of territorial males and number of interactions 
between conspecific males (for 20 min in each h) in the main territory centering on map 
coordinates 070100. Intermittent line, number of interactions. Number of males in ter- 
ritory: line I, 1; line II, 2; line III, 8. Bottom: Frequency of territorial males and patrollers 


VOLUME 41, NUMBER 1 55 


appeared, and males patrolled as much in the afternoon as in the 
morning (perch: patrol, 1:2.75 to 1:2; x?,) = 0). 


Territories 


Location of territories. Transect data over the common revealed that 
males establish territories at edge or junction sites mainly at woodland 
margins. Sites without nectar are less favorable as males establish ter- 
ritories in these areas only in the afternoon. A comparison of two groups 
each comprising four woodland margins equal in size and similar in 
aspect differed in the number of territories they contained on the basis 
of a nectar source, bramble (x?) = 6.72, P < 0.01). 

Repeated transects (6/h) in the clearing throughout the day for sev- 
eral days revealed that territories were established in two main areas 
(Fig. 5): in the middle of the clearing (map grid reference 070100) and 
in the NW part of the site (map grid reference 1222). The canopy edge 
facing direct sunlight, the tall birch seedlings in the center of the 
clearing and the main body of heather and furze were all generally 
ignored. Crowding of the clearing by males led to a number of subsid- 
iary territories being established in the afternoon, but the main terri- 
tories occupied edge sites, exposed to sunshine, between the hostplant 
and some other vegetation type. The central territory was not affected 
by shade at any time of the day and included an extensive area of E. 
tetralix, a major nectar source. By comparison, the NW territory was 
abandoned in the late afternoon when it became shaded and nectar 
sources were limited. 

Except for the central territory, the pattern of perches was noticeably 
affected by the diurnal changes in shade, a significant shift in territories 
occurring from W to E as the day progressed (x?) = 16.38, P < 0.001). 
None of the perch sites below vertical coordinate 07 was occupied before 
1200 h. Apart from this diurnal shift, the pattern of territories remained 
virtually stable from year to year. There is, however, clear indication 
of a seasonal shift in perches in the central territory, a significant 
movement northwards in perches between 8 and 7 July 1984 (x?) = 
7.41, P < 0.01) and between 7 and 23 July 1984 (x, = 6.38, P < 0.02). 
This coincided with a shift in flowering of E. tetralix, from the shorter 
Molinia zone to the Calluna sward where it is partially overtopped. 
Males were perching 10 cm higher on 23 July and therefore in a cooler 
microclimate during cooler conditions (maximum temperatures: 7 July, 
27°C; 23 July, 24°C; windspeed: 7 July, 3.4 knots; 23 July, 5.9 knots). 


— 

from 48 transects covering clearing. Mean hourly number is obtained by dividing numbers 
in activity by 6. The few males feeding have been omitted. Shade temperatures are given 
for each hour (°C). 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


11.12 13° 4hs 159 dee 


1031213) 6) 15. aaa 





Fic. 5. Probability distribution map for territorial male O. venata, based on 162 
transects in clear conditions evenly spaced over 4 days, 30 June, 3, 7, and 23 July 1984 
(6 transects/h). Two well established territories occur; the most important centers on map 
coordinate 0710, the second around 1422. 


VOLUME 41, NUMBER 1 57 





1 


Fic. 6. Fish-net plot illustrating relative frequency of different perch sites by territorial 
male O. venata in main territory of clearing (map grid reference 070100) based on 38 
insects and 364 records (resolution 10 cm). The figures at the corners are map coordinates 
for Fig. 1. 


Inside territories. There is a distinction between what can be labelled 
a territory (the area defended by each male) and perches within a 
territory (the platforms or sites used by males after each sortie or 
voluntary patrol in the area defended). Dominant among perches in 
the main territory of the clearing are tiny birch seedlings under 15 cm 
tall (the same height or lower than surrounding heather), corresponding 
to the six substantial peaks in the plot (Fig. 6). The multiplicity of 
residual “‘relief’’ relates to sprigs of heather or cross-leaved heath, even 
grass blades, casually used during voluntary patrols or after interactions. 

The actual size and shape of the territory varies from individual to 
individual (Fig. 7) despite the fidelity of different insects from day to 
day and year to year to the main perches illustrated in Fig. 6. Many 
males are restricted to some smaller portion of the two territories in 
the clearing; in the central territory, to the E or W half separated by 
the bank of Calluna. Nevertheless, others use a wider array of perches, 
and the unified structure of the main territory is evident in the links 
joining the main perches in Fig. 7. Interactions between males lead to 
the discovery and use by incumbents of a wider array of perches. Thus, 
both the main territories in the clearing are too large for one O. venata; 
intruders can pass over a portion of the territory and temporarily settle 
in another part of it unnoticed. However, each territory is also too small 
for two or more O. venata. A sortie after intruders by one occupant 
usually triggers activity in the other. Once this happens, violent inter- 
actions occur, often repeatedly triggered by a third party, until one of 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


N 





3 4 5 6 7 8 9 10 11 


Fic. 7. Linkage diagram illustrating consecutive moves by male O. venata between 
perch sites in main territory of clearing around grid coordinate 0710. Zone covered by 
peaks in Fig. 6 represents a single territorial unit. Frequency of moves between perches 
is not illustrated. Data as in Fig. 6. Base map includes location of vegetation boundaries 
and tree seedlings (squares) shown in Fig. 1. 


the two prior occupants leaves. Occasionally such premium is placed 
on the central territory that fourth and fifth males were noticed to enter 
while three contesting the zone are immersed in dispute. They, in turn, 
become incorporated in the next “dog fight”. 

The siting of territories has priority over perches, although shape and 
size of a territory can be distorted by altering the nature of perches 
within it. Males occasionally perched on artificial objects on the com- 
mon, and used these repeatedly even if these “perches’’ were moved 
about. This effect is shown for the clearing where Fig. 8 illustrates perch 
sites adopted with and without a white plastic bag, alternated 4 times 
over 10 min periods. Clearly, the plastic bag disrupts the pattern of 
activity (Fig. 8, x24) = 51.6, P < 0.001) and acts as an overoptimal 


—_ 

Fic. 8. Linkage diagrams illustrating consecutive moves by male O. venata between 
perch sites in main territory of clearing; Top: in the presence of an artificial object (white 
plastic shopping bag). Bottom: in its absence. Triangles show two locations used for the 
artificial perch. 


VOLUME 41, NUMBER 1 





60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


stimulus compared to the territory. There is, however, a limit to which 
territorial sites can be distorted. Artificial perches are used in the vicinity 
of sites where territories would normally be established, but apparently 
not in open areas usually ignored by the butterfly. At the woodland 
edge, the plastic bag was occupied continuously over 17 min of obser- 
vation and contested for 13 times. In an adjacent wet heath area, it was 
visited twice by male O. venata during 11 min who inspected it while 
in flight and moved on. 


DISCUSSION 


O. venata males reveal similarities and some differences with other 
butterflies that should complement the discussion on mate location 
behavior. Pertinent questions are: What determines the distribution of 
males? Why do they both perch and patrol? Why do they change their 
activities through the day, and how do they know when to change? 
Does the butterfly engage in territorial defense and, if it does, what 
factors induce the butterfly to defend one area more vigorously than 
another? How does it recognize good territorial sites and suitable perch- 
es in them? 

Males are absent from the woodland cover and the dry heath. The 
woodland cover is dense, little light reaches the ground, and the but- 
terflies would not be able to thermoregulate under it. The other negative 
zone, the dry heath, is void of nectar sources and lacks larval hostplants. 
Males are found in Molinia areas where there are eclosion sites, the 
most predictable locations for females. The enormous bias for bramble 
bushes points to a combination of factors: bramble is a major nectar 
source, and Molinia occurs in the immediate vicinity. Moreover bram- 
ble is typically associated with edge and junction sites. 

O. venata perches and establishes territories in the early morning 
when air temperatures are low, and during the afternoon when they 
are highest. However, the insect treats afternoon as morning if the early 
part of the day has been cloudy and cool. In this respect, it is similar 
to vanessids in behavior, which also establish territories in the afternoon 
(Baker 1972, Bitzer & Shaw 1979, 1983), but differ from three satyrids, 
L. megera, P. aegeria, and C. pamphilus, which patrol more when air 
temperatures are highest (Dennis 1982a, Shreeve 1984, Wickman 1985a). 
Adoption of perching implies that a particular resource is more difficult 
to obtain. For one or more of several reasons, scrambling for the resource 
over a wider area becomes ineffective, and it is then necessary to wait 
for females at vantage points. The reasons are: (i) males have insufficient 
energy supplies to remain patrolling; more specifically, energy used in 
patrolling begins to exceed that used in defense (Baker 1972). (ii) Am- 
bient conditions are inadequate for sustained flight, which lowers body 


VOLUME 41, NUMBER 1 61 


temperatures (Shreeve 1984). (iii) The ratio of available females is 
substantially reduced, and patrolling becomes less effective than perch- 
ing at vantage points, regardless of energy resources. Energy losses 
during perching may match those of patrolling as skirmishing in ter- 
ritories can often be continuous and more violent than patrolling flights. 
Some territories are also occupied in the mornings, during the early 
and late flying season and in prime locations, presumably when energy 
levels are high. 

Early morning perching may well relate, in part, to the need for 
both sexes to warm up and become fully active, but does not explain 
afternoon perching and mid-morning patrolling. On clear days, tem- 
peratures are higher after 1800 h than before. Perching as opposed to 
patrolling is more likely to relate to the unavailability of unmated 
females. It is possible that most unpaired females may be found by 
patrolling in the early part of the day, but that a switch to perching is 
effected in the afternoon as patrolling for a dwindling resource becomes 
unsuccessful. Whatever the explanation for switches in behavior, the 
reason is closely tied to the cue used by males to determine when 
switches are made. As neither energy nor ambient conditions (heat and 
sunshine) seem sufficient to explain the timing of switches in behavior, 
contacts (or lack of them) and scent may form prominent cues. The 
way males are arrested in flight to scan small areas for several minutes 
where nothing is to be seen suggests this. Clearly, more needs to be 
known about the influence of scent, particularly the distances over 
which behavior can be influenced by it. 

Perching (and in some cases patrolling [Shields 1967]) has been likened 
to territoriality by Baker (1972) since defense of some “‘resource’’ is 
involved. This is denied by Scott (1974) for three reasons: (i) Males are 
incapable of learning topographic details of sites, and move on to new 
areas. (ii) As obvious resources are missing, there can be no territories 
for feeding, roosting, or oviposition. (iii) Interactions involve investi- 
gation of sex and not defense. Baker (1972), Davies (1978) and Dennis 
(1982a) have shown that males return to the same area repeatedly, even 
to the same perches, and that butterflies are capable of a spatial learning 
process (Baker 1978). We agree that species are genetically imprinted 
for characteristic topographic sites in which to establish territories (Scott 
1974:107), but not for the precise details of sites, as insinuated by Scott. 
Scott’s insistence that males should occupy the same area during several 
days does not hold in any case, since occupancy should be gauged 
against the time required to fulfill a particular function. Some perches 
are clearly established where there are female resources (hostplant sites 
in A. urticae [Baker 1972]; possible thermoregulation sites in P. aeqgeria 
[Parker 1978]; oviposition, therefore emergence, sites in L. megera [Den- 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


nis 1982a, 1986]). In O. venata, males have a bias for establishing ter- 
ritories where there is nectar in addition to the hostplant. Even where 
fundamental resources for females are missing, researchers have in 
‘every case shown that perches correspond to strong linear features or 
prominent visual markers, which have the capacity for concentrating 
resources. For virgin females, the most essential resource is males. Such 
strong visual lines dictate movement in insects and determine predict- 
able locations for both sexes (Baker 1972, Dennis 1982a, Dennis & 
Bramley 1985, Shields 1967, Bitzer & Shaw 1979, 1983). Scott’s own 
research has shown this to be the case in hilltopping males (Scott 1968) 
and for perching skippers (Scott 1973). 

Finally, Baker (1972, 1983) has indeed shown that defense can be 
involved, and his observations are substantiated by others (Davies 1978, 
Dennis 1982a, Bitzer & Shaw 1979, 19838, Wickman & Wiklund 1983). 
In O. venata, perch sites are undoubtedly defended and are thus ter- 
ritories. Baker’s (1972, 1983) observations are confirmed in that, usually, 
intruders failed to settle in an area unless the area was vacated, and 
unless escalated contests occurred when ownership was disputed. Two 
issues point away from Scott’s insistence that territoriality in butterflies 
is no more than the investigation of the sex of the intruder by the 
incumbent and avoidance by the intruder of a would-be predator. First, 
interactions are significantly shorter between patrolling males; clearly 
it does not take long for males to determine the sex of individuals. 
Interactions between males at perch sites involve escalated contests back 
and forth over the perch site. Secondly, if the intruder is attempting 
to escape from a predator, it is then difficult to explain why it returns, 
often repeatedly, to the site to invite further strikes from the “‘predator’’. 
Despite these criticisms, it would be a mistake to regard the territorial 
defense (Baker 1983) and investigation (Scott 1974) models as mutually 
incompatible, although Scott’s reasons for denying territorial defense 
are wrong. We suspect that the scale of defense is as variable as mate 
location behavior is flexible. At least three features point to an ability 
in O. venata to assess costs and benefits in defense: (i) the degree to 
which occupancy and defense varied between different territories in 
the clearing (corresponding to the premium placed on sites by males 
measured in numbers of intruders, length and violence of interactions); 
(ii) the varying degrees of pugnacity between contacting males when 
engaged in different activities (feeding, patrolling, and perching); and 
(iii) the voluntary abandonment of territories after unsuccessful periods. 

Territories are typically established along distinct linear features, 
visual markers such as edges and junctions, but there is some indication 
that female resources may also be a factor in the location (as well as 
degrees of defense) of territories. Some sites, such as those in the clearing 


VOLUME 41, NUMBER 1 63 


(Fig. 5) associated with nectar sources, had territories throughout the 
morning on many days, but other sites along the woodland fringe 
associated with hostplant but without nectar were established only in 
the afternoon. Moreover, there was a distinct shift in location of the 
main territory in the clearing (Fig. 5) which coincided with changes 
in nectar apparency but not with temperatures. The varying frequency 
of perches in different parts of this clearing is in itself evidence of the 
varying quality of sites. However, not all hostplant locations and emer- 
gence sites become territories. Open areas, where females were found 
eclosing, were ignored. Visual rather than scent cues seem to be prom- 
inent in setting up territories. 

Perching spots in territories are repeatedly used after interactions, 
but the fidelity to particular perches depended on the array of oppor- 
tunities (number of potential perches) and the degree of disturbance 
by intruders, which usually led incumbents to cover a wider area and 
take up different posts. Typical perches were low, robust launching 
platforms providing effective observation posts. Taller seedlings over 1 
m high were avoided, presumably because incumbents would then not 
be able to pick out intruders against the background vegetation beneath 
them. O. venata learns the spatial configuration of its territory quickly. 
Searches for artificial objects used as perches, removed during inter- 
actions, are first made where they occurred, the insects thereafter in- 
creasing the area searched. Voluntary patrols within the area of the 
territory probably contribute to gaining familiarity with territory land- 
marks. Several features then combine to demonstrate the adaptability 
of this hesperiid: opportunist behavior, spatial memory, ability to rec- 
ognize resource-generating landmarks, and perhaps location of mates 
using scent—a developing theme in butterflies demonstrating degrees 
of “intelligence” in short-lived temperate species as well as tropical 
relatives with long-lived adults. 


ACKNOWLEDGMENTS 


We thank T. G. Shreeve for many helpful comments. Fish-net plots were produced 
using SURFACE-2 graphics via the Durham University AMDAHL 470V8 computer. 


LITERATURE CITED 


BAKER, R. R. 1972. Territorial behavior of the nymphalid butterflies, Aglais urticae 
(L.) and Inachis io (L.). J. Anim. Ecol. 41:453-469. 

1978. The evolutionary ecology of animal migration. Hodder and Stoughton. 

1012 pp. 

1983. Insect territoriality. Ann. Rev. Entomol. 28:65-89. 

BELLINGER, P. F. 1954. Attraction of zebra males by female pupae. J. Lepid. Soc. 8: 
102. 

BINK, F. A. 1985. Hostplant preferences of some grass feeding butterflies. Proc. 3rd 
Congr. Eur. Lepid. 1982. Pp. 23-29. 








64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BiTzeER, R. J. & K. C. SHAW. 1979. Territorial behavior of the red admiral, Vanessa 
atalanta (L.) (Lep: Nymphalidae). J. Res. Lepid. 18:36-49. 

1983. Territorial behavior of Nymphalis antiopa and Polygonia comma. J. 
Lepid. Soc. 37:1-13. 

COURTNEY, S. P. 1980. Studies in the biology of the butterflies Anthocharis cardamines 

(L.) and Pieris napi (L.) in relation to speciation in Pierinae. Ph.D. Thesis, Durham. 
244 pp. 

DAVIES, N. B. 1978. Territorial defense in the speckled wood butterfly, Pararge aegeria: 
The resident always wins. Anim. Behav. 26:138-147. 

DENNIS, R. L. H. 1982a. Mate location strategies in the wall brown butterfly, Lasiom- 
mata megera L. (Lep: Satyridae). Wait or seek? Entomol. Rec. J. Var. 94:209-214; 
95:7-10. 

1982b. Patrolling behavior in orange tip butterflies within the Bollin valley in 

north Cheshire, and a comparison with other pierids. Vasculum 67:17-25. 

1986. Motorways and cross-movements. An insect’s “mental map’ of the M56 
in Cheshire. Amat. Entomol. Soc. Bull. (in press). 

DENNIs, R. L. H. & M. L. BRAMLEY. 1985. The influence of man and climate on 
dispersion patterns within a population of adult Lasiommata megera (L.) (Satyridae) 
at Brereton Heath, Cheshire. Nota Lepid. 8:309-324. 

HEATH, J., E. POLLARD & J. A. THOMAS. 1984. Atlas of butterflies in Britain and Ireland. 
Viking. 158 pp. 

PARKER, G. A. 1978. Evolution of competitive mate searching. Ann. Rev. Entomol. 23: 
173-196. 

PEACHEY, C. A. 1980. The ecology of the butterfly community of Bernwood Forest. 
M. Phil. Thesis, Oxford Polytechnic. 151 pp. 

POLLARD, E. 1977. A method for assessing changes in the abundance of butterflies. Biol. 
Conserv. 12:115-134. 

1979. A national scheme for monitoring the abundance of butterflies: The first 
three years. Proc. Brit. Entomol. Nat. Hist. Soc. 12:77-90. 

RuTOWSKI, R. L. 1983. Sexual selection and the evolution of butterfly mating behavior. 
J. Res. Lepid. 23:125-142. 

ScoTT, J. A. 1968. Hilltopping as a mating mechanism to aid survival of low density 
species. J. Res. Lepid. 7:191—204. 

1972. Mating of butterflies. J. Res. Lepid. 11:99-127. 

1973. Adult behavior and population biology of two skippers mating in con- 

trasting topographic sites. J. Res. Lepid. 12:181-196. 

1974. Mate-locating behavior in butterflies. Am. Midl. Nat. 91:103-117. 

1975. Mate-locating behavior of western North American butterflies. J. Res. 

Lepid. 14:1—40. 

1983. Mate-locating behavior of western North American butterflies. II. New 
observations and morphological adaptations. J. Res. Lepid. 21:177-187. 

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6:69-178. 

SHREEVE, T. G. 1984. Habitat selection, mate location and microclimatic constraints 
on the activity of the speckled wood butterfly, Pararge aegeria. Oikos 42:371-877. 

WICKMAN, P.-O. 1985a. The influence of temperature on the territorial and mate 
locating behaviour of the small heath butterfly, Coenonympha pamphilus (L.) (Lep- 
idoptera: Satyridae). Behav. Ecol. Sociobiol. 16:233-238. 

1985b. Territorial defense and mating success in males of the small heath 
butterfly, Coenonympha pamphilus L. (Lepidoptera: Satyridae). Anim. Behav. 33: 
1162-1168. 

WICKMAN, P.-O. & C. WIKLUND. 1983. Territorial defense and its seasonal decline in 
the speckled wood (Pararge aegeria). Anim. Behav. 31:1206-1216. 
































Received for publication 11 June 1986; accepted 5 November 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 65-69 


THE STATUS OF “PAPILIO HIPPARCHUS” STAUDINGER 
(PAPILIONIDAE) 


KURT JOHNSON 


Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024 


AND 


DAVID MATUSIK 


Department of Entomology, Field Museum of Natural History, 
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605 


ABSTRACT. The holotype male and only known specimen of “P. hipparchus”’ is 
critically examined for the first time since its description a century ago. Although tra- 
ditionally accorded species status solely on the basis of Staudinger’s original description 
and figure, this specimen in fact represents a morph of Protesilaus phaon Boisduval. 

Additional key words: Protesilaus phaon, Colombia, taxonomy. 


“Papilio hipparchus”’ Staudinger (1884) (type locality Cauca [Colom- 
bia]) has been one of the most confusing members of the “lysithous- 
related group” of swallowtail butterflies. Munroe (1961) placed this 
group in a subgenus of Eurytides Hiibner. Hancock (1983) accorded 
the group generic status as Protesilaus Swainson. Irrespective of this 
difference, both authors included the following taxa: asius (Fabricius), 
microdamas (Burmeister), thymbraeus (Boisduval), belesis (Bates), 
branchus (Doubleday), ilus (Fabricius), lysithous (Hiibner), ariarathes 
(Esper), harmodius (Staudinger), trapeza (Rothschild & Jordan), xynias 
(Hewitson), phaon (Boisduval), euryleon (Hewitson), pausanias (Hew- 
itson), protodamas (Godart), hipparchus (Staudinger), kumbachi (Vo- 
geler), and chibcha (Fassl). It has since been demonstrated that illu- 
minatus (Niepelt), dospassosi (Riitimeyer) and huanucana (Varea 
deLuque) also belong to this group (Johnson et al. 1986a, 1986b). 

Protesilaus hipparchus has been traditionally accorded species status 
solely on the basis of Staudinger’s original description and figure of the 
type. This holotype (in the Staudinger collection at the Zoologisches 
Museum der Humbolt Universitat zu Berlin [ZMH]) has not been ex- 
amined by twentieth century students of Papilionidae (Rothschild & 
Jordan 1906, Jordan 1907, D’Almeida 1965, D’Abrera 1981, Hancock 
1983). Based on original descriptions, Hancock (pers. comm.) speculated 
that P. hipparchus, P. chibcha and P. kumbachi represent aberrations. 

As part of our ongoing review of some papilionid groups, and as aid 
to colleagues preparing a synonymic list of South American Papilion- 
idae, we obtained the type (male) of P. hipparchus for study. It is 
described below, and dorsal and ventral surfaces, attached labels, and 
Staudinger’s original figure are illustrated (Fig. 1) as well as relevant 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 











Spee “on. 
ee oy) 






e h. Au rchus 
i doe 






Origin) aie 
| 


Fic. 1. Recent photograph of holotype male Papilio hipparchus Staudinger. A, Upper- 
surface; B, Under-surface. C, Photograph of Staudinger’s (1888) painted figure of P. 
hipparchus showing upper-surface on left, and under-surface on right. 


genitalic characters (Fig. 2). Genitalic characters of Protesilaus have 
been reviewed in detail elsewhere (Johnson et al. 1986a, 1986b). As 
noted by these and other studies (Munroe 1961, Hancock 1983, Beu- 
talspacher & Howe 1984), the valval harpe provides the most diagnostic 
characters. 


The Papilio hipparchus Type 
(Figs. 1A, B, 2A) 


Length of forewing (base to apex): 40.5 mm. 

Upper-surface of wings: Ground blackish brown. Forewing submarginal markings gray, 
hued slightly yellowish; hindwing submarginal markings gray, hued slightly yellowish, 
medial band very dull gray-white (faintly tinged with pink, a trait which might not be 
considered worthy of mention had it not been emphasized in the original description and 
subsequent interpretations of authors). 

Under-surface of wings: Ground blackish brown. Forewing submarginal markings 
gray-white, hued slightly yellowish caudad; hindwing submarginal markings gray-white, 
hued slightly yellowish, a slight reddish slash in each cell costad to M1 and basad to each 
submarginal marking; medial band a slight lightening of ground color, becoming more 
obsolescent costad to M2 (Fig. 1B and C exaggerate extent of this-lightness); two anal 


VOLUME 41, NUMBER 1 67 





Fic. 2. Diagnostic features of male genitalia of P. hipparchus holotype (A), and 
representative male of P. phaon phaon (Colombia, AMNH), after Johnson et al. 1986a, 
1986b (B). Each shows inner-lateral view of right valve. 


markings yellowish distad, reddish centrad. Traces of red occur at base of both wings 
along the thorax. 
Genitalia (Fig. 2A): Differing negligibly from nominate P. phaon (Fig. 2B). 


DISCUSSION 


The type of P. hipparchus does not represent a valid species, but 
rather, a morph of P. phaon. The genitalia are indistinguishable from 
P. phaon. P. phaon is highly variable as shown by the number of infra- 
specific names proposed for it (Rothschild & Jordan 1907:661-663, 
D’Abrera 1981:62). Early workers suggested that P. hipparchus should 
be associated with P. euryleon, probably as a sister species (Rothschild 
& Jordan 1906, Jordan 1907). This view probably resulted from Stau- 
dinger’s original description. There he states that the accompanying 
figure is not accurate in all details, and that P. hipparchus lacks par- 
ticular wing markings of P. euryleon, a species belonging to a mono- 
phyletic group that includes P. phaon, P. pausanias, P. protodamas, 
and P. illuminatus (Johnson et al. 1986a, 1986b, K. S. Brown, pers. 
comm.). Jordan (1907) and Rothschild and Jordan (1906) speculated 
that the inaccuracies of the original figure involved (a) the frequency 
of red-spotting in the anal area of the hindwing under-surface, and (b) 
the extent of the under-surface medial band. The degree of these mark- 
ings could constitute major differences between the wing patterns of 
P. hipparchus and P. euryleon. However, as indicated in our description 
above, (a) the red on the under-surface is indeed nearly absent, and 
(b) the medial band (shown in the Staudinger figure as a brown band 
proceeding costad to the discal cell [Fig. 1C]) is actually a simple light- 
ening of the under-surface ground color extending across the entire 
wing. In both of these features, the type of P. hipparchus resembles 
morphs of P. phaon more than those of P. euryleon. The major error 
in the original figure concerns the extent of submarginal markings on 
the forewing upper-surface. While the original figure shows these ex- 
tending only slightly costad (Fig. 1C), they actually extend costad to 
the apex, where they are darker caudad. 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


We do not know whether this morph represents a natural population 
of possible subspecific status or a one-time occurrence. Were it not for 
slight yellowish tinges to submarginal wing markings, and the almost 
imperceptible pinkish flush to the upper-surface medial hindwing band, 
P. hipparchus might be considered a “‘black-white” morph of Prote- 
silaus. Such black-and-white phenotypes are reported to occur as mim- 
ics of black-and-white-marked papilionids of the tribe Troidini (Young 
1971, K. S. Brown, pers. comm.). They include such Protesilaus as P. 
illuminatus Niepelt (Johnson et al. 1986b), P. harmodius female form 
viginia Rothschild & Jordan (D’Abrera 1981), P. phaon male form 
ulopos Gray, and a tentative subspecies of P. euryleon from near Buga 
in the Cauca Valley of Colombia. Such mimicry probably also explains 
the unique black-white morph of recently described Heraclides ma- 
tusiki Johnson & Rozycki (1986) (Papilionidae, Papilionini). Interpre- 
tation of P. hipparchus as not a black-white morph may follow only 
from the emphasis on red and yellowish markings in the original de- 
scription, and its elaboration by subsequent authors without access to 
the type. There is no way to know if the type specimen has faded, but 
our experience indicates fading is unlikely. The type of Staudinger’s 
“Papilio diaphora”’ (Johnson et al. 1985) is just as old, and suggests no 
fading when compared with recent specimens. 


ACKNOWLEDGMENTS 


Review comments by D. L. Hancock (National Museum, Republic of Zimbabwe, 
Bulawayo, Zimbabwe), P. R. Ackery (British Museum, Natural History) and an anonymous 
reviewer were helpful. We thank H. J. Hannemann (ZMH) for loan of the type specimen 
and K. S. Brown Jr. (Universidade Estadual de Campinas, Sao Paulo, Brazil) for help 
with species-level taxa in the Papilionidae. The following persons searched collections 
and answered queries: P. R. Ackery, Rienk de Jong (Leiden, Netherlands), D. L. Hancock, 
O. H. H. Mielke (Curitaba, Brazil), L. D. Miller (Sarasota, Florida), Tommasso Racheli 
(Rome, Italy), J. E. Rawlins (Pittsburgh, Pennsylvania), R. K. Robbins (Washington, D.C.), 
Richard Vane-Wright (London, England), E. W. Schmidt-Mumm (Caracas, Venezuela). 


LITERATURE CITED 


BEUTALSPACHER, C. R. & W.H. HOWE. 1984. Mariposas de Mexico. Fasiculo I., Papilion- 
idae. La Preusa Medica Mexicana, S.A., Mexico City. xii + 128 pp. 

D’ABRERA, B. 1981. Butterflies of the neotropical realm. Part 1. Papilionidae and Pier- 
idae. Landsdowne Editions, East Melbourne. 172 pp. 

D’ALMEIDA, R. F. 1965. Catalogo dos Papilionidae americanos. Sociedade Brasileria de 
Entomologia, Sao Paulo. 366 pp. 

HANCOCK, D. 1983. Classification of the Papilionidae (Lepidoptera): A phylogenetic 
approach. Smithersia 2:1—48. 

JOHNSON, K., D. MATUsIK & R. ROZYCKI. 1986a. A study of Protesilaus microdamas 
(Burmeister) and the little-known P. dospassosi (Riitimeyer) and P. huanucana (Varea 
de Luque) (Papilionidae). J. Res. Lepid. 25 (in press). 

JOHNSON, K. & R. ROZYCKI. 1986. A new species of the anchisiades group of Heraclides 
from Venezuela (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 94:383-393. 


VOLUME 41, NUMBER 1 69 


JOHNSON, K., R. Rozyck! & D. MATusIK. 1985. Species status and the hitherto unrec- 
ognized male of Papilio diaphora Staudinger (1981), (Lepidoptera: Papilionidae). J. 
New York Entomol. Soc. 93:99-109. 

1986b. Rediscovery and species status of the neotropical swallowtail butterfly 
Papilio illuminatus Niepelt (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 
94:516-525. 

JORDAN, K. 1907. Papilionidae, pp. 1-51. In Seitz, A. (ed.), Macrolepidoptera of the 
world. Vol. 5. Alfred Kernen Verlag, Stuttgart. vii + 592 pp. 

MUNROE, E. 1961. The classification of the Papilionidae (Lepidoptera). Can. Entomol. 
Suppl. 17. 51 pp. 

ROTHSCHILD, W. & K. JORDAN. 1906. A revision of the American papilios. Novit. Zool. 
13:412-752. 

STAUDINGER, O. 1884[1888]. Exotische Tagfalter, Theil I. In Staudinger, O. & E. Schatz 
(eds.), Exotische Schmetterlinge. Band 1, Beschreibungen, 333 pp.; Band 2, Abbil- 
dungen, 100 plts. 

YOuNG, A. M. 1971. Mimetic associations in natural populations of tropical papilionid 
butterflies (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 79:210-224. 





Received for publication 9 June 1986; accepted 29 October 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 70-74 


A NEW WHITE-AND-BLACK SUBSPECIES OF 
PROTESILAUS EURYLEON (PAPILIONIDAE) 


KURT JOHNSON 


Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024 


AND 


DAVID MATUSIK 


Department of Entomology, Field Museum of Natural History, 
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605 


ABSTRACT. Protesilaus euryleon pleiades (tribe Leptocircini) is described from the 
southern Cauca Valley of Colombia. The uniqueness of its completely white-and-black 
wing pattern is discussed in relation to other recently described taxa of Protesilaus and 
Heraclides (Papilionidae, tribe Papilionini) which apparently mimic white-and-black 
papilionids of the tribe Troidini. 


Additional key words: Protesilaus euryleon pleiades, taxonomy, Colombia. 


Attention has been drawn to neotropical papilionids of the tribes 
Papilionini and Leptocircini (sensu Hancock 1983) which display wing 
patterns primarily white over darker ground color (Johnson et al. 1986a, 
1986b). Such phenotypes, distinctive among taxa usually showing red 
or orange-red hindwing markings, are generally attributed to mimicry 
of white-and-black colored taxa of the papilionid tribe Troidini (Johnson 
et al. 1986a, 1986b, Young 1971, K. S. Brown, pers. comm.). Our 
examination of examples of these white and dark mimetic morphs has 
led to increased recognition of their occurrence. Heraclides matusiki 
Johnson & Rozycki (1986) was described from NE Venezuela, and 
represents a distinctive cream-white-and-black taxon in the anchisiades 
species group (sensu Munroe 1961, Hancock 1988) (tribe Papilionini). 
Protesilaus illuminatus (Niepelt) (tribe Leptocircini), formerly known 
from one extant male syntype and an accompanying female (and not 
mentioned in the literature since its 1928 description) was collected 
again in 1981 and studied by us (Johnson et al. 1896b). This taxon, 
which we accorded species status because of its cream-white-and-black 
wing markings and distinctive genitalia, requires biological study to 
ascertain its relation to red-and-black morphs of the remaining species 
of the group. P. illuminatus might represent a biological subspecies of 
either P. euryleon (Hewitson) or P. ariarathes (Esper). In the P. eu- 
ryleon-related species cluster of Protesilaus (which includes P. illu- 
minatus, P. euryleon, P. phaon (Boisduval), P. pausanias (Hewitson), 
and P. protodamas (Godart) [Johnson et al. 1986a, 1986b, K. S. Brown, 
pers. comm.]) only the P. phaon male form ulopos (Gray) has been 


VOLUME 41, NUMBER 1 7A 


recognized as a primarily white-and-black mimetic morph (D’Abre- 
ra 1981). In remaining Protesilaus, P. harmodius xenaides female form 
virginia (Rothschild & Jordan) is an example (D’Abrera 1981), and 
possibly also the type and only specimen of P. hipparchus (Staudinger). 
We have shown that the latter taxon represents a morph of P. phaon 
(Johnson & Matusik 1987). 

During our study of the above papilionid groups, we obtained from 
the Calima River region of the southern Cauca Valley of Colombia 
fresh specimens of a completely white-and-black morph of P. euryleon 
(Fig. 1). The specimens are genitalically indistinguishable from P. eu- 
ryleon (Fig. 2), but their wing patterns of pristine white on velvetine 
black differ from all known P. euryleon populations. 

P. euryleon is exceedingly polymorphic (D’Abrera 1981:62-68, 
Rothschild & Jordan 1906:663-666). However, all previously known 
populations of the species show red coloration of the hindwing orbs, 
and most have prominent white to yellowish (or occasionally greenish) 
patches located medially on both surfaces of the forewing. 

The value of the subspecies concept has been a source of controversy 
among lepidopterists, particularly regarding nearctic taxa (Murphy & 
Ehrlich 1984). In the neotropics, however, where mimicry phenomena 
abound (Sheppard et al. 1985), the concept has particular heuristic 
value and utility. Therefore, we apply the subspecies category to the 
recently discovered white-and-black southern Cauca Valley population 
of P. euryleon. 


Protesilaus euryleon pleiades, new subspecies 


(Figs. 1A, B, 2A) 


Diagnosis. Distinguishable from all congeneric species-level taxa except P. illuminatus 
by the completely white-and-black wing markings except for minor reddish colorations 
as noted below. P. illuminatus has triangular-shaped white hindwing markings, not orb- 
shaped as in P. euryleon. P. e. pleiades has submarginal white coloration typical of the 
species, and lacks the prominent dorsal red anal marking of P. illuminatus. P. e. pleiades 
is distinct from other infraspecific white-and-black morphs of Protesilaus as follows: P. 
h. xenaides female form virginia (as characteristic of harmodius) has a complete medial 
line of small round spots on the hindwing upper- and under-surfaces (but white in form 
virginia); P. phaon male form ulopos (as characteristic of phaon) lacks the whitish medial 
patch on both forewing surfaces. 

Description. Male. Upper-surfaces of wings: Ground color velvetine black. Forewing 
with bright white medial patch; hindwing with bright white orbs in vein interspaces from 
anal margin to caudad of vein M2; light white along margin at vein interspaces, more 
emphatic costad and bordered basad by extremely narrow red slashes in limbal area. 
Under-surface of wings: Ground color velvetine black. Forewing with area of mimetic 
patch powdered blackish over vague white; hindwing with markings as on upper-surface 
but slightly duller; however, red slashes are more emphatic basad of marginal spots and 
distinct at anal margin. Length of forewing: 41.0 mm (holotype); 40.5 mm (paratype). 
Female unknown. Male genitalia (Fig. 2A): Typical of P. euryleon as described by Johnson 


72 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fic. 1. Holotype male of P. e. pleiades. A. Upper-surface; B. Under-surface. 


and Rozycki (1986) and Johnson et al. (1986b), differing only in more sharply pointed 
keel of valval harpe, and slightly larger ventral process (Fig. 2). 

Types. Holotype male, on hillside above Calima River, 1,500 m altitude, 50 km SW 
of Buga, Cauca Valley, Colombia, 15 June 1983, leg. Phillip Mays, in American Museum 
of Natural History (AMNH); paratype male, same data, leg. local collector with P. Mays, 
in collection of Phillip Mays (Tarzana, California); paratype male, Rio Bravo, Calima 
River Valley [in Cauca Valley], June 1985, leg. Charles Condor, in collection of Rick 
Rozycki (Chicago, Illinois). 

Distribution. Presently known only from the type locality, but possibly of wider dis- 
tribution. 

Remarks. Because we received the primary type and first-listed paratype specimens 
second-hand through an anonymous commercial dealer (who mentioned three other male 
specimens and provided conflicting collection data on different occasions) there was initial 
confusion about the precise location of the P. e. pleiades population. Earlier, K. S. Brown 
Jr. informed us he had heard reports of the population but did not know its whereabouts 
or the whereabouts of any specimens. Details were provided when we learned from Mays 
that he had personally participated in collecting the eventual primary type and one 
paratype. We assume the three other reported males (currently in the hands of the above- 
mentioned anonymous dealer) come from the same general area because Mays visited 
the type locality after being told that three other specimens had been collected there. 
Mays received this information from Christopher Farrell, a dealer who formerly resided 
in Colombia. Farrell reportedly remembered the general collecting data on the three 
specimens, though he did not keep them. The Cauca Valley data is further supported by 
the recent collection of the second-listed paratype. Considering the above, earlier data 
given us (and perhaps others) by the commercial source, citing the vicinity of Leticia, 
Amazonas State, Colombia, for P. e. pleiades specimens is probably inaccurate. Since 
these data derived from verbal communication, with label data available only in the case 
of the Mays paratype, we think the Leticia data were a miscommunication. P. illuminatus 
occurs N of Leticia in the upper Rio Putumayo Valley, and had also been obtained by 
us through the same commercial source; this may have caused the confusion. This back- 
ground deserves mention because a white-and-black form of P. euryleon could foreseeably 
occur in the upper Rio Putumayo region, at the edge of its generally montane range. 


VOLUME 41, NUMBER 1 73 





Fic. 2. Male genitalia of (A) P. e. euryleon, Costa Rica, AMNH, after Johnson et al. 
(1986a, 1986b) and (B) P. e. pleiades holotype. Each shows inner-lateral view of right 
valve. 


Another papilionid known only from this region, P. dospassosi (Riitimeyer), also is dis- 
tinctive in its reduction of upper surface red coloration (Johnson et al. 1986a). 

Etymology. Foliowing on the Greek binomial and referring to the bright white-on- 
black markings, pleiades (from the constellation of that name) denotes the seven-spotted 
pattern of the wing surfaces. 


ACKNOWLEDGMENTS 


Review comments by T. C. Emmel (University of Florida, Gainesville) and an anon- 
ymous reviewer were helpful. We thank the latter for detailed suggestions concerning 
distributional data. Phillip Mays kindly provided important details. K. S. Brown Jr. 
(Universidade Estadual de Campinas, Sao Paulo, Brazil) discussed subject material with 
us and he and the following persons searched collections or answered queries: P. R. Ackery 
(London, England), Rienk de Jong (Leiden, Netherlands), D. L. Hancock (Bulawayo, 
Zimbabwe), O. H. H. Mielke (Curitaba, Brazil), L. D. Miller (Sarasota, Florida), Tommasso 
Racheli (Rome, Italy), J. E. Rawlins (Pittsburgh, Pennsylvania), R. K. Robbins (Wash- 
ington, D.C.), Richard Vane-Wright (London, England), and E. W. Schmidt-Mumm 
(Caracas, Venezuela). 


LITERATURE CITED 


D’ABRERA, B. 1981. Butterflies of the neotropical realm. Part 1. Papilionidae and Pier- 
idae. Landsdowne Editions, East Melbourne. 172 pp. 

Hancock, D. 1988. Classification of the Papilionidae (Lepidoptera): A phylogenetic 
approach. Smithersia 2:1—48. 

JOHNSON, K. & D. Matusik. 1987. The status of “Papilio hipparchus” Staudinger 
(Papilionidae). J. Lepid. Soc. 41:65-69. 

JOHNSON, K., D. MATuSIK & R. ROZYCKI. 1986a. A study of Protesilaus microdamas 
(Burmeister) and the little-known P. dospassosi (Rutimeyer) and P. huanucana (Varea 
de Luque) (Papilionidae). J. Res. Lepid. 25 (in press). 

JOHNSON, K. & R. ROZYCKI. 1986. A new species of the anchisiades group of Heraclides 
from Venezuela (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 94:383-393. 

JOHNSON, K., R. Rozycki & D. Matusik. 1986b. Rediscovery and species status of the 
neotropical swallowtail butterfly Papilio illuminatus Niepelt (Lepidoptera: Papilion- 
idae). J. New York Entomol. Soc. 94:516-525. 

MUNROE, E. 1961. The classification of the Papilionidae (Lepidoptera). Can. Entomol. 
Suppl. 17. 51 pp. 

Murpny, D. D. & P. R. EHRLICH. 1984. On butterfly taxonomy. J. Res. Lepid. 23: 
19-34. 

ROTHSCHILD, W. & K. JORDAN. 1906. A revision of the American papilios. Novit. Zool. 
13:412-752. 


74 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


SHEPPARD, P. M., J. R. G. TURNER, K. S. BROWN, W. W. BENSON & M. C. SINGER. 1985. 
Genetics and the evolution of Muellerian mimicry in Heliconius butterflies. Phil. 
Trans. R. Lond., B 308:433-610. 

YOUNG, A. M. 1971. Mimetic associations in natural populations of tropical papilionid 
butterflies (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 79:210-224. 


Received for publication 9 June 1986; accepted 30 October 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 75-76 


GENERAL NOTE 


PREDATION ON ADULTS OF ANARTIA FATIMA (FAB.) 
Additional key words: Nymphalidae, Costa Rica. 


Vertebrate predation on adults can be a significant source of mortality in butterfly 
populations (Bowers et al. 1985, Evolution 39:93-103), yet the incidence of such predation 
has been documented in few cases. Although direct observations of vertebrate predation 
are rare (exceptions: Brown & Vasconcellos-Neto 1976, Biotropica 8:136-141; Fink & 
Brower 1981, Nature 291:67-70; Ehrlich & Ehrlich 1982, J. Lepid. Soc. 37:148-152), 
several studies have examined museum specimens or specimens collected during field 
sampling for evidence of bird predation (Carpenter 1941, Proc. Zool. Soc. Lond. A 1941: 
223-231; Shapiro 1974, Am. Nat. 108:229-232, and others). Characteristic damage in- 
flicted by birds (and probably lizards) includes symmetrical tears on the wings, straight 
cuts across major veins, triangular tears, and beak imprints (Sargent 1973, J. Lepid. Soc. 
27:175-192; Bowers & Wiernasz 1979, Ecol. Entomol. 4:205-209). In general, unpalatable 
butterfly species have a higher incidence of beak imprints due to birds tasting the butterfly 
or remembering a previous bad experience and voluntarily releasing it (Shapiro, cited 
above; Bowers & Wiernasz, cited above). In contrast, palatable butterflies have a higher 
incidence of beak tears due to ripping their wings out of the bird’s beak (Shapiro, cited 
above; Bowers & Wiernasz, cited above). 

This study assessed the incidence of predation on adults of the common, palatable 
(Silberglied et al. 1980, Science 209:617-619) butterfly, Anartia fatima (Fab.) (Nym- 
phalidae), at Finca La Selva Biological Station, Costa Rica. 

Sixty A. fatima adults were collected in the grassy area near the laboratory at Finca 
La Selva and at “Rafael’s house” by the river on 8 March 1986. The butterflies were 
stored in envelopes and later sexed and assessed for evidence of predation. Butterflies 
showing potential signs of predation were examined for characteristic indications of bird 
or lizard attack (Bowers & Wiernasz, cited above). 

The sex ratio of our sample of 60 butterflies was 38 males and 22 females (1.7:1). Seven 
of the 60 butterflies (12%) showed clear evidence of predation, 6/38 males (16%), and 
1/22 females (4.5%). Although there were more males damaged than females, the dif- 
ference was not significant (Fisher Exact Test, P, = 0.38, power [1 - 6] = 0.21 [Zar 1984, 
Biostatistical analysis, 2nd ed., Prentice Hall, New Jersey]). Four individuals had sym- 
metrical damage on two hindwings (HW) only, and another showed damage on two 
hindwings and a forewing (FW), indicating that the individuals were attacked while the 
wings were held together. Two individuals showed damage on a single side, one on the 
right HW-FW, and the other on one FW, indicating that the butterflies were probably 
attacked while the wings were open (Bowers & Wiernasz, cited above) such as when 
basking or flying. These results suggest that most predation occurred while the butterflies 
were roosting, probably at dawn or dusk (Rawlins & Lederhouse 1978, J. Lepid. Soc. 32: 
145-159). 

Twelve % predation is similar to what has been found in some other butterfly populations 
such as Cercyonis pegala (Nymphalidae: Satyrinae) (Bowers & Wiernasz, cited above) 
and Ascia monuste (Pieridae) (Pough & Brower 1977, Am. Mid]. Natur. 98:50-58). We 
were unable to distinguish damage potentially caused by birds or lizards, and both may 
prey on butterflies (Boyden 1976, Evolution 30:73-81; Ehrlich & Ehrlich, cited above; 
refs. in Bowers et al., cited above). 

The damaged individuals we collected were those that escaped after being attacked. 
We found only beak tears on the wings, and no beak imprints, concordant with the known 
palatability of these butterflies to predators (refs. in Silberglied et al. 1979, Psyche 87: 
219-260; Harrison & Crabtree, pers. obs.). Our small sample indicates that individuals 
attacked at the hindwings are more likely to escape, perhaps because of the fragility of 
these wings compared to the forewings. Anartia fatima, as a palatable butterfly, thus 
likely escapes by tearing its wings out of the attacking animal’s mouth or beak. This would 
be an easy task if the butterfly were captured by the flimsy hindwings. 


76 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Results from this study, as well as that of Silberglied et al. (1980, above) indicate that 
predation on these butterflies is relatively common. Although we do not have information 
on the number of successful attacks by predators on A. fatima, its palatability to both 
vertebrate and invertebrate predators coupled with the incidence of damage suggests 
that such predation may be a significant source of mortality. 

This project was supported by funds from the Organization of Tropical Studies. 


M. DEANE BOWERS, Museum of Comparative Zoology, Harvard University, Cam- 
bridge, Massachusetts 02138. 

ROSE C. CRABTREE, Biological Laboratories, Harvard University, Cambridge, Mas- 
sachusetts 02138. 

SUSAN P. HARRISON, Department of Entomology, University of California, Davis, 
California 95616. 

CLAUDIA SOBREVILLA, Harvard University Herbaria, Harvard University, Cambridge, 
Massachusetts 02138. 

MICHAEL WELLS, School of Forestry and Environmental Studies, Yale University, 
New Haven, Connecticut 06511. 

Lorne M. WOLFE, Department of Ecology, Ethology, and Evolution, University of 
Illinois, Urbana, Illinois 61820. 


Received for publication 27 May 1986; accepted 28 October 1986. 


Journal of the Lepidopterists’ Society 
41(1), 1987, 77-81 


OBITUARY 


JOHN STEVEN BUCKETT (1939-1986) 


John Steven Buckett was born 6 December 1939 in Oakland, California, and died of 
pneumonia in Davis, California, 16 January 1986. 

Steve attended primary and secondary schools in Petaluma, California, and graduated 
from Petaluma High School in 1957. Following brief service in the United States Marine 
Corps, he enrolled in Santa Rosa Junior College in 1958. 

Steve entered the University of California, Davis, in September 1960. He earned his 
Bachelor of Science with a major in entomology in January 1965 and a Master of Science 
in entomology one year later. While a student at Davis, Steve became a member of Phi 
Kappa Phi in 1966 and Sigma Xi in 1967. 

Between 1967 and 1969 Steve was employed as a systematic entomologist with the 
California Department of Agriculture in Sacramento. He was a Life Member of the 
Lepidopterists’ Society and served briefly as Treasurer in 1969. 

Steve wished to complete his Ph.D. at Davis but deteriorating health prevented this 
goal. His interest in moths remained intense, and he continued to collect until 1984, even 
though his activities became restricted to his yard in Davis. Although his fingers were 
crippled by psoriasis, he still prepared specimens occasionally during most of 1985. 

The door to Steve's house was never locked. Friends and visitors were always welcome, 
and especially so when conversation turned to natural history and, particularly, moths. 

Steve is survived by his mother, Mrs. Bertha R. Buckett, and sister, Mrs. Patricia Moore, 
both living in Santa Rosa, California. 

Although Steve had broad interests in the natural sciences, his many publications dealt 
mostly with millipeds and moths. The joint moth collections of William Bauer and Steve 
Buckett, and the extensive Buckett library, have been donated to the R. M. Bohart 
Museum, Department of Entomology at the University of California, Davis. 





John Steven Buckett, 1963 


78 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NEW TAXA DESCRIBED BY JOHN STEVEN BUCKETT 


The holotypes of these species are deposited in the R. M. Bohart Museum, Department 
of Entomology, University of California, Davis, California 95616, unless otherwise in- 
dicated. 


Lepidoptera—Geometridae 


Philtraea utahensis Buckett 1970 [1971]. J. Res. Lepid. 9:41. American Museum of 
Natural History. 

Philtraea surcaliforniae Buckett 1970 [1971]. J. Res. Lepid. 9:43. 

Philtraea latifoliae Buckett 1970 [1971]. J. Res. Lepid. 9:47. 

Philtraea mexicana Buckett 1970 [1971]. J. Res. Lepid. 9:53. United States National 
Museum of Natural History. 

Philtraea albimaxima Buckett 1970 [1971]. J. Res. Lepid. 9:55. Los Angeles County 
Museum of Natural History. 

Philtraea monillata Buckett 1970 [1971]. J. Res. Lepid. 9:62. American Museum of 
Natural History. 


Lepidoptera—Noctuidae 


Abagrotis kirkwoodi Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 
14:8. 

Abagrotis rubricundis Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 
15:11. 

Abagrotis striata Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 15:18. 

Abagrotis glenni Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 15:14. 

Abagrotis hennei Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 15:16. 

Abagrotis tecatensis Buckett 1968. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 
16:16. 

Abagrotis reedi Buckett 1969. Calif. Dept. Agric. Bureau Entomol. Occ. Pap. No. 17:6. 

Anepia plumasata Buckett & Bauer 1967. J. Lepid. Soc. 21:235. 

Annaphila macfarlandi Buckett & Bauer 1964. J. Res. Lepid. 3:99. 

Behrensia conchiformis suffusa Buckett 1964 [1965]. J. Res. Lepid. 3:189. Type deposi- 
tory unknown. 

Euxoa piniae Buckett & Bauer 1964. Can. Entomol. 96:967. 

Faronta terrapictalis Buckett 1967 [1969]. J. Res. Lepid. 6:268. 

Feralia meadowsi Buckett 1967 [1968]. J. Res. Lepid. 6:43. 

Nephelodes adusta Buckett 1972 [1973]. J. Res. Lepid. 11:260. 

Oncocnemis sandaraca Buckett & Bauer 1966 [1967]. J. Res. Lepid. 5:197. 

Petaluma Buckett & Bauer 1964 [1965]. J. Res. Lepid. 3:193. 

Petaluma californica Buckett & Bauer 1964 [1965]. J. Res. Lepid. 3:194. 

Petalumaria Buckett & Bauer 1967 [1968]. J. Res. Lepid. 6:52. 

Polia piniae Buckett & Bauer 1966 [1967]. J. Res. Lepid. 5:221. 

Pseudocopivaleria Buckett & Bauer 1966. J. Lepid. Soc. 20:84. 

Pseudocopivaleria anaverta Buckett & Bauer 1966. J. Lepid. Soc. 20:88. 

Xylomiges baueri Buckett 1967 [1968]. J. Res. Lepid. 6:25. 


Diplopoda—Idagonidae 


Idagonidae Buckett & Gardner 1967. Mich. Entomol. 1:117. 
Idagona Buckett & Gardner 1967. Mich. Entomol. 1:120. 
Idagona westcotti Buckett & Gardner 1967. Mich. Entomol. 1:120. 


Diplopoda—Lysiopetalidae 

Tynomma magnum Buckett & Gardner 1969. Pan-Pac. Entomol. 45:207. 
Diplopoda—Spirobolidae 

Tylobolus deses magnificus Buckett & Gardner 1966. Proc. Biol. Soc. Wash. 79:45. 


VOLUME 41, NUMBER 1 79 


Diplopoda—Vanhoeffeniidae 


Bidentogon Buckett & Gardner 1968. Pan-Pac. Entomol. 44:198. 
Bidentogon helferorum Buckett & Gardner 1968. Pan-Pac. Entomol. 44:198. 


Diplopoda—X ystodesmidae 


Anombrocheir Buckett & Gardner 1969. Entomol. News 80:69. 

Anombrocheir spinosa Buckett & Gardner 1969. Entomol. News 80:70. 

Anombrocheir bifurcata Gardner & Buckett 1969. Entomol. News 80:295. 

Harpaphe haydeniana lanceolata Buckett & Gardner 1968. Calif. Dept. Agric. Bureau 
Entomol. Occ. Pap. No. 11:22. 

Harpaphe haydeniana maurogona Buckett & Gardner 1968. Calif. Dept. Agric. Bureau 
Entomol. Occ. Pap. No. 11:23. 

Metaxycheir Buckett & Gardner 1969. Proc. Entomol. Soc. Wash. 71:67. 

Metaxycheir prolata Buckett & Gardner 1969. Proc, Entomol. Soc. Wash. 71:67. 

Wamokia discordalis Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:522. 

Wamokia hoffmani Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:528. 

Wamokia dentata Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:533. 

Wamokia falcata Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:534. 

Wamokia remota Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:535. 

Wamokia sierrae Buckett & Gardner 1968. Proc. Biol. Soc. Wash. 81:537. 


BIBLIOGRAPHY OF JOHN STEVEN BUCKETT 


BUCKETT, J.S. 1968. Collecting of Annaphila spila with notes on the “Crimson-winged” 
group of the genus. J. Res. Lepid. 2:303-304. 

1964. Rediscovery and redescription of the moth, Euxoa marinensis (Lepidop- 

tera: Noctuidae). Pan-Pac. Entomol. 40:101-104. 

1964. Identity of the moth, Euxoa wilsoni (Lepidoptera: Noctuidae). Pan-Pac. 

Entomol. 40:104-107. 

1964. Annotated list of the Diplopoda of California. Simmons Publishing Com- 

pany, Davis, CA, 34 pp. 

1964. Distributional notes on Septis maxima with illustrations of the genitalia 

(Lepidoptera: Noctuidae). Pan-Pac. Entomol. 40:162-164. 

1964 [1965]. Revision of the North American genus Behrensia with a description 

of a new subspecies. J. Res. Lepid. 3:129-144. 

1964. Review of the Depicta group of the genus Annaphila with the description 

of a new species from Oregon. J. Res. Lepid. 3:95-101. 

1964 [1965]. Petaluma, a new genus, with the description of a new species. J. 

Res. Lepid. 3:193-196. 

1965 [1966]. The noctuid moth Annaphila baueri with notes on its habits. J. 

Res. Lepid. 4:185-189. 

1965 [1966]. A reevaluation of Annaphila casta (Noctuidae). J. Res. Lepid. 

4:199-204. 

1965. Identity of Heliosea celeris melicleptroides with notes on its habits. J. 

Res. Lepid. 4:79-80. 

1966. Distributional patterns of certain members of the noctuid genus Annaphila 

(Lepidoptera: Noctuidae). Bull. Entomol. Soc. Am. 12:113. 

1966. Distributional notes on Apamea acera (Smith) with discussion of the type 

specimen (Lepidoptera: Noctuidae). Pan-Pac. Entomol. 42:283-286. 

1966 [1967]. Rediscovery of Annaphila casta Hy. Edw. in California (Noctui- 

dae). J. Res. Lepid. 5:37-388. 

1966. The mantid Stagmomantis limbata (Hahn) in California (Orthoptera: 

Mantidae). Pan-Pac. Entomol. 42:57-58. 

1966. Key to the genera of Psaphidini, with descriptions of a new genus and 

species from western North America (Noctuidae: Cuculliinae). J. Lepid. Soc. 20: 

83-91. 















































80 










































































JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1966 [1967]. The little known moth Euxoa sculptilis (Harvey) in Arizona, with 
descriptions, illustrations, and notes on Euxoa violaris (Grote and Robinson) (Noc- 
tuidae—Agrotiinae). J. Res. Lepid. 5:255-261. 

1966 [1967]. Discovery of a larval host plant for Annaphila lithosina with notes 
on the species (Noctuidae: Amphipyrinae). J. Res. Lepid. 5:262-264. 

1966 [1967]. A new species of Oncocnemis from the western United States 
(Noctuidae: Cuculliinae). J. Res. Lepid. 5:197-208. 

1966 [1967]. A new species of Polia Ochsenheimer from California and notes 
on Polia discalis (Grote) (Noctuidae: Hadeninae). J. Res. Lepid. 5:221-228. 

1967. Agrotid moths of the southern nearctic region. Am. Philos. Soc. 1967: 
255-256. 

1967 [1968]. Description of a new species of Xylomiges from California, with 
notes and illustrations (Lepidoptera: Noctuidae: Hadeninae). J. Res. Lepid. 6:23-30. 
1967 [1968]. A new species of Feralia from Santa Catalina Island of California, 
with notes on the immature stages of Feralia februlis Grote (Noctuidae: Cuculliinae). 
J. Res. Lepid. 6:438-51. 

1967 [1969]. A new species of armyworm. J. Res. Lepid. 6:268-274. 

1967 [1968]. Homonymy of the “new genus’ Petaluma Buckett & Bauer (Noc- 
tuidae) under Petaluma Hulst (Pyralidae) and proposal of the name Petalumaria 
for “Petaluma” californica. J. Res. Lepid. 6:52. 

1967. Description of a new species of Anepia Hampson from the Sierra Nevada 
of California (Noctuidae). J. Lepid. Soc. 21:285-240. 

1967. A new family of cavernicolus millipeds with the description of a new 
genus and species from Idaho (Diplopoda: Chordeumida: Chordeumidea). Mich. 
Entomol. 1:117-126. 

1968. Rediscovery of the type of the milliped, Harpaphe telodonta (Chamberlin) 
(Diplopoda: Xystodesmidae). J. New York Entomol. Soc. 76:60-63. 

1968. Description of the male of Lithophane gausapata (Noctuidae). J. Lepid. 
Soc. 22:42—45. 

1968. Revision of the Nearctic moth genus Abagrotis Smith with descriptions 
of new species (Lepidoptera: Noctuidae). Calif. Dept. Agric. Bureau Entomol. Occ. 
Pap. No. 12:1-21. 

1968. Revision of the Nearctic moth genus Abagrotis Smith with descriptions 
of new species (Lepidoptera: Noctuidae). Calif. Dept. Agric. Bureau Entomol. Occ. 
Pap. No. 14:1-16. 

1968. Revision of the Nearctic moth genus Abagrotis Smith with descriptions 
of new species (Lepidoptera: Noctuidae). Calif. Dept. Agric. Bureau Entomol. Occ. 
Pap. No. 15:1-29. 

1968 [1969]. Identity of the moth “Stretchia” behrensiana (Grote) with new 
synonymy (Noctuidae). J. Res. Lepid. 7:57-68. 

1968. A new genus and species of milliped from northern California (Polydes- 
mida: Vanhoeffeniidae). Pan-Pac. Entomol. 44:198-202. 

1968. Revision of the milliped genus Wamokia Chamberlin from the Sierra 
Nevada of central California (Diplopoda: Polydesmida: Xystodesmidae). Proc. Biol. 
Soc. Wash. 81:511-538. 

1968. Revision of the milliped genus Harpaphe Cook from western North 
America (Polydesmida: Xystodesmidae). Calif. Dept. Agric. Bureau Entomol. Occ. 
Pap. No. 11:1-51. 

1968 [1970]. Species in the genera Polia Ochsenheimer and Euxoa Hitibner from 
the western United States (Lepidoptera: Noctuidae). J. Res. Lepid. 7:87-94. 

1968 [1970]. Revision of the Nearctic moth genus Abagrotis Smith with de- 
scriptions of new species (Lepidoptera: Noctuidae). Calif. Dept. Agric. Bureau Ento- 
mol. Occ. Pap. No. 16:1-27. 

1969. A new genus of xystodesmid milliped from northern California. Entomol. 
News 80:67-78. 

1969. A new genus of xystodesmid milliped with description of a new species 
from Idaho (Diplopoda: Xystodesmidae). Proc. Entomol. Soc. Wash. 71:65-70. 


VOLUME 41, NUMBER 1 81 





1969. A review of the genus Gosodesmus Chamberlin, with the synonomy of 
Encybe Chamberlin (Diplopoda: Polydesmida: Andrognathidae). J. New York Ento- 
mol. Soc. 76:40-50. 

1969. Revision of the chordeumid milliped genus Tynomma Loomis from 

California (Lysiopetalidea: Lysiopetalidae). Pan-Pac. Entomol. 45:204-216. 

1969. Revision of the Nearctic moth genus Abagrotis Smith with descriptions 

of new species (Lepidoptera: Noctuidae). Calif. Dept. Agric. Bureau Entomol. Occ. 

Pap. No. 17:3-27, 238 figs. 

1969 [1970]. Identity of the moth Loxagrotis pampolycala (Dyar) from the 

southwestern United States and Mexico (Noctuidae). J. Res. Lepid. 8:118-128. 

1970 [1971]. Revision of the Nearctic genus Philtraea Hulst, with notes on 

biology and the descriptions of new species (Geometridae). J. Res. Lepid. 9:29-64. 

1971 [1973]. Identity of the moth Oncocnemis semicollaris J. B. Smith, with 

notes and distribution. J. Res. Lepid. 10:248-254. 

1972 [1973]. A new species of Nephelodes Guenée from the Great Basin area 
of northeastern California (Noctuidae: Hadeninae). J. Res. Lepid. 11:260-268. 
BUCKETT, J. S. & W. R. BAUER. 1964. A new species of Euxoa Hbn. (Lepidoptera: 

Noctuidae) from the Sierra Nevada of California. Can. Entomol. 96:967-970. 

BUCKETT, J. S. & R. H. LEUSCHNER. 1965 [1966]. Rediscovery and redescription of the 
moth Lithophane vanduzeei (Barnes), with notes on the type locality (Lepidoptera: 
Noctuidae). J. Res. Lepid. 4:281-286. 

BUCKETT, J. S. & L. P. Lounrpos. 1965 [1966]. The little known species Luperina 
venosa. A redescription of the species with additional distributional data. J. Res. 
Lepid. 4:227-232. 

BUCKETT, J. S. & M. R. GARDNER. 1966. Rediscovery and redescription of Tylobolus 
deses Cook, with the description of a new subspecies (Diplopoda: Spirobolidae). Proc. 
Biol. Soc. Wash. 79:41-48. 

BUCKETT, J. S. & T. A. SEARS. 1968 [1970]. Variation in color and maculation in a 
population of Nemoria pulcherrima from the Sierra Nevada of California (Lepi- 
doptera: Geometridae). J. Res. Lepid. 7:95-98. 

OPLER, P. A. & J. S. BUCKETT. 1970 [1971]. Seasonal distributional of “Macrolepidop- 
tera” in Santa Clara County, California. J. Res. Lepid. 9:75-88. 




















ROBERT O. SCHUSTER, R. M. Bohart Museum of Entomology, Department of Ento- 
mology, University of California, Davis, California 95616, AND JULIAN P. DONAHUE, 
Los Angeles County Museum of Natural History, 900 Exposition Boulevard, Los An- 
geles, California 90007. 


Date of Issue (Vol. 41, No. 1): 16 April 1987 


Ul eat A Run pony 











' eo , as ok a 
‘ie j pei J. mle) WUSIt iit y \ 

: J s : vy 
g = ' 

at 


r ie 


li ey ii: ¢ omerlieet) ait F dikegih , 


“hy” Ln!) Comet epee Gh tae 
“> Ca cobra My, At Weuptan 


Aol sp av’, 6 PSR 


. “Ma Aw fr 4 
vibe} } 


~ 
+8 
“| 
{ 4 
: na i 
1 ‘ 1 a) 








EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
ROBERT C. LEDERHOUSE, THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Categories of contributions are Articles, General Notes, Technical Com- 
ments, Book Reviews, and Obituaries. Short manuscripts concerning new state records, 
current events, and notices should be sent to the News, June Preston, Editor, 832 Sunset 
Drive, Lawrence, KS 66044 U.S.A. Journal contributors should prepare manuscripts 
according to the following instructions. 

Abstract: An informative rather than descriptive abstract should precede the text of 
Articles. 

Key Words: Up to five key words or terms not in the title should accompany Articles, 
General Notes, and Technical Comments. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely 
double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. 
Titles should be explicit and descriptive of content, including the family name of the 
subject, but must be as short as possible. The first mention of a plant or animal in the 
text should include the full scientific name with author. Measurements should be given 
in metric units; times should be given in terms of the 24-hour clock (0930 h, not 9:30 
AM). Underline only where italics are intended. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In General Notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30). 

Illustrations: Only half of symmetrical objects such as adults with wings spread should 
be illustrated, unless whole illustration is crucial. Photographs and drawings should be 
mounted on stiff, white backing, arranged in the desired format, allowing (with particular 
regard to lettering) for reduction to their final width (usually 4% inches). Illustrations 
larger than 8% x 11 inches are not acceptable and should be reduced photographically 
to that size or smaller. The author’s name, figure numbers as cited in the text, and an 
indication of the article’s title should be printed on the back of each mounted plate. 
Figures, both line drawings and halftones (photographs), should be numbered consecu- 
tively in Arabic numerals. The term “plate” should not be employed. Figure legends 
must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), 
headed EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of 
illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the proofs. 
A page charge of $60 per full Journal page is applicable to all authors. Unfunded authors 
may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A 





CONTENTS 


STATUS AND HABITATS OF POTENTIALLY ENDANGERED LEPI- 
DOPTERA IN OHIO. John A. Shuey, Eric H. Metzler, David 
C. Iftner, John V. Calhoun, John W. Peacock, Reed A. Wat- 
kins, Jeffrey D. Hooper & William F. Baboock cccccccumene 


MILKWEED PATCH QUALITY, ADULT POPULATION STRUCTURE, AND 
ECG LAYING IN THE MONARCH BUTTERFLY. M. P. Zalucki 
d> Y. Suzuki 20 ON 


HOST SPECIFICITY AND BIOLOGY OF BUCCULATRIX IVELLA BUSCK, 
A POTENTIAL BIOLOGICAL CONTROL AGENT FOR BACCHARIS 
HALIMIFOLIA L. IN AUSTRALIA. W. A. Palmer & G. 
Diatloff 00 ee 

BUTTERFLIES FROM THE UJUNG KULON NATIONAL PARK, 
INDONESIA. T.R. New, M. B. Bush & H. K. Sudarman ... 


THE IDENTITY OF CYCLOPIDES PAOLA PLOTZ (HESPERIIDAE). 
David L; Hancock \00000 oY 


MATE LOCATION BEHAVIOR OF THE LARGE SKIPPER BUTTERFLY 
OCHLODES VENATA: FLEXIBLE STRATEGIES AND SPATIAL 
COMPONENTS. R. L. H. Dennis & W. R. Williams _........ 


THE STATUS OF “PAPILIO HIPPARCHUS’ STAUDINGER (PAPILIO- 
NIDAE). Kurt Johnson’ G David Matusik ae 


A NEW WHITE-AND-BLACK SUBSPECIES OF PROTESILAUS EURYLEON 
(PAPILIONIDAE). Kurt Johnson & David Matusik 0. 


GENERAL NOTE 


Predation on adults of Anartia fatima (Fab.). M. Deane Bowers, Rose C. 
Crabtree, Susan P. Harrison, Claudia Sobrevilla, Michael Wells & Lorne 
M. Wolfe: 2 EO he 


ANNOUNCEMENT 
Technical Comments: a new category in the Jourmal cee 


OBITUARY 
John Steven Buckett (1989-1986) 0 


13 


23 


29 


4] 


45 


65 


Volume 41 3 1987 Number 2 


ISSN 0024-0966 





JOURNAL 


of the 


y 
cue 


_ Leprpoprerists’ Sociery 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





26 June 1987 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


DoucLas C. FERGUSON, President GERARDO Lamas M., Vice President 
Jerry A. POWELL, President-Elect EBBE SCHMIDT NIELSEN, Vice 
CLIFFORD D. FERRIS, Immediate Past President President 

OLAF H. H. MIELKE, Vice President JAMES P. TUTTLE, Treasurer 


RICHARD A. ARNOLD, Secretary 


Members at large: 


BoYcE A. DRUMMOND III MIRNA M. CASAGRANDE M. DEANE BOWERS 
JOHN LANE EDWARD C. KNUDSON RICHARD L. BROWN 
ROBERT K. ROBBINS FREDERICK W. STEHR PAUL A. OPLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists' Society, to: James P. Tuttle, Treasurer, 
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian 
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007- 
4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Secretary, 
50 Cleaveland Rd., #3, Pleasant Hill, CA 94523-3765, U.S.A. 


To obtain: 

Back issues of the Journal and NEWS (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society”) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, CA 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los 
Angeles, CA 90007-4057. Second-class postage paid at Los Angeles, CA and additional 
mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ Society, % 
Natural History Museum, 900 Exposition Blvd., Los Angeles, CA 90007-4057. 


Cover illustration: Semilooping larva of the strange noctuid Phyprosopus callitrichoides 
on Smilax. Sketch by Mark Klingler, Carnegie Museum of Natural History. Suggested 
by John E. Rawlins. 


JouRNAL OF 
Tue LeEpPIpDOPTERISTS’ SOCIETY 


Volume 41 1987 Number 2 


Journal of the Lepidopterists’ Society 
41(2), 1987, 83-93 


ECOLOGICAL SIGNIFICANCE OF A POSTMATING 
DECLINE IN EGG VIABILITY IN THE 
TIGER SWALLOWTAIL 


ROBERT C. LEDERHOUSE AND J. MARK SCRIBER! 


Department of Entomology, University of Wisconsin, 
Madison, Wisconsin 53706 


ABSTRACT. The number of times that field-collected tiger swallowtail females had 
mated increased significantly with age. Mean number of spermatophores per female was 
1.97, and 65% of the females had mated more than once. Egg fertility and viability were 
independent of the number of eggs a female had oviposited. Both parameters declined 
significantly with time for sequential samples of eggs from both hand-paired and field- 
collected females. Between 10% and 25% of field-collected females that carried a sper- 
matophore and laid over 15 eggs produced no viable eggs. Fresh singly-mated females 
and worn multiply-mated females did not differ in egg viability, but singly-mated worn 
females had significantly lower egg viabilities. Thus, an additional mating may be ad- 
vantageous for females that receive inadequate spermatophores or live for more than a 
week. 


Additional key words: Papilionidae, Papilio glaucus, infertility, spermatophore counts, 
multiple matings. 


Mating histories of female tiger swallowtails, Papilio glaucus L. (Pa- 
pilionidae), as revealed by spermatophore counts, probably are docu- 
mented better than for any other species of butterfly (Drummond 1984). 
Although the debate about the role of sexual selection in maintaining 
the female color polymorphism remains unresolved (Burns 1966, Pliske 
1972, Platt et al. 1984), it is clear that multiple mating is common in 
this species. This is in contrast to suggestions on theoretical grounds 
that female butterflies should prefer monogamy (Wiklund 1977a, 1977b, 
1982). Levin (1973) was unable to demonstrate experimentally a selec- 
tive advantage for multiple mating in P. glaucus. 

The maintenance of the color polymorphism in female tiger swal- 


1 Present address of both authors: Department of Entomology, Michigan State University, East Lansing, Michigan 
48824. 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lowtails requires frequency-dependent selection or a balancing of se- 
lection pressures (Burns 1966, 1967, Prout 1967, Levin 1973). The loci 
that control it are on the sex chromosomes (Clarke & Sheppard 1962, 
Scriber 1985). Dark-morph females benefit as Batesian mimics of the 
unpalatable, aposematic pipevine swallowtail, Battus philenor (L.) 
(Brower 1958). One subspecies, P. glaucus canadensis, which occurs N 
of the range of B. philenor, has only monomorphic yellow females. 
Both yellow and dark female morphs occur in the other two subspecies, 
P. g. glaucus and P. g. australis, which are sympatric with the model, 
and the relative frequency of the dark morph is correlated positively 
with the abundance of the model (Brower & Brower 1962, Lederhouse 
& Scriber 1987). 

Burns (1966) documented a greater mean number of spermatophores 
carried by field-collected yellow females than by dark females. He 
suggested that sexual selection favored yellow females by shortening 
the interval between emergence and mating. Yellow females would 
also benefit if a single spermatophore were inadequate to fertilize all 
eggs a female was likely to oviposit. However, subsequent spermato- 
phore-count studies of other P. glaucus populations have not revealed 
significant differences in spermatophore number for the two color morphs 
(Pliske 1972, 1973, Makielski 1972, Platt et al. 1984). In addition, Levin 
(1973) showed that a single spermatophore was sufficient to fertilize all 
the eggs laid by P. glaucus females in the laboratory. This eliminated 
preferential mating with respect to color as the balancing selective 
pressure, but did not explain why roughly half of the females in these 
studies mated more than once. 

However, closer examination of Levin’s data revealed that his ex- 
perimental females averaged only about 118 eggs during 5.4 days. Both 
Remington (1959) and Lederhouse (1981) observed declines with time 
in the viability of eggs from singly mated females of other swallowtail 
species. In the black swallowtail, P. polyxenes F., a decline in fertility 
was detectable only after about 10 days and was independent of the 
number of eggs laid (Lederhouse 1981). In the present study, our pur- 
pose was to investigate the fertility and viability of eggs laid by P. 
glaucus females with longer mating intervals than those in the study 
by Levin (1978). In addition, we wanted to determine under what 
circumstances a female would benefit from multiple matings. 


MATERIALS AND METHODS 


Samples of females were collected from several locations. The P. g. 
glaucus used for determining number of spermatophores carried were 
captured in Adams and Scioto counties, Ohio. The P. g. canadensis 
were collected in several counties in north-central Wisconsin. These 


VOLUME 41, NUMBER 2 85 


females were assigned to one of four categories of wing wear when 
they were collected (Lederhouse 1978). After they died, field-collected 
females and a portion of hand-paired females were dissected, and the 
number of spermatophores was determined. 

Laboratory-reared (lab) females were hand-paired once to field-col- 
lected (field) or laboratory-reared males. All females were set up in 
plastic boxes (10 cm xX 20 cm X 27 cm) with a sprig of black cherry, 
Prunus serotina, under saturated humidity. The boxes were placed 
about 70 cm from continuously lighted 100-watt incandescent bulbs. 
Females were fed a mixture of 1 part honey to 4 parts water at least 
once daily. Most females were allowed to oviposit until death. 

Eggs were collected and counted at two-day intervals except on 
weekends. Larvae were removed as they hatched, and the remaining 
eggs were monitored for 10 days after the last larva hatched. Because 
the green eggs of P. glaucus become mottled and then turn black as 
the embryo develops, eggs could be assigned to one of three mutually 
exclusive categories. Eggs were classified as infertile if they showed no 
development, inviable if they developed but did not hatch, or viable 
if they hatched. 

Unless otherwise indicated, values are presented as means + standard 
deviations. Since the lab-reared males and females were the offspring 
of mothers from a variety of locations, and since numerous combinations 
were mated, the Friedman two-way analysis of variance was used for 
the analyses of fertility and viability (Siegel 1956). Sequential samples 
from each female were ranked for both variables. A modified Tukey 
test was used for multiple comparisons of ranked samples (Zar 1984). 
Only females that laid eggs for at least three sampling intervals, pro- 
duced at least 50 eggs, and had some eggs hatch were included in these 
analyses. In addition, each field-collected female with at least one sper- 
matophore and that laid 15 or more eggs was monitored for egg fertility 
and viability. 


RESULTS 


Field-collected dark-morph females averaged 1.97 + 0.90 sper- 
matophores (Table 1). All had mated at least once, and 65.4% had 
mated more than once. The females were distributed uniformly across 
wear classes with a mean of 2.45. Mean number of spermatophores per 
female increased significantly with increasing wing wear (Table 1). The 
correlation between spermatophore number and wear class was highly 
significant (Spearman Rank Correlation, r = 0.52, P < 0.001). 

Although the actual percent hatching was highly variable from fe- 
male to female (Table 2), the pattern of decreasing egg viability was 
detectable by the end of the first week of oviposition. All laboratory- 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Number of spermatophores contained by P. glaucus females collected in 
Adams Co., Ohio in relation to wing condition. z and P values are for one-tailed Mann- 
Whitney U-tests comparing the means for adjacent condition classes. 


Number of spermatophores 


Condition n ] 2 3 4 Mean Z P 





Fresh 23 16 6 1 0 1.35 2.30 <0.02 
Slightly worn 34 13 a 2 2 1.79 , 

: 1.82 <0.04 
Intermediate 29 7 12 9 1 2.14 2.18 <0.02 
Very worn 21 1 8 8 4 7,7 Ah 

Total 107 37 43 20 d 1.97 





reared groups of females showed a significant decline for the first three 
samples of eggs (Friedman two-way analysis of variance, Table 38). 
When only females that laid eggs for four intervals were considered, 
again all groups of lab females had significant decreases in viability 
(Table 4). The reverse pattern occurred in the analysis of infertility. 
The only group without a significant infertility increase through three 
samples was the 1984 lab females paired with lab males (Table 3). All 
three groups of lab females had significant increases in infertility through 
four samples (Table 4). There was no discernible pattern for the inviable 
category of eggs, although one group had a significantly nonhomoge- 
neous result. In sum, the decline in viability with sample resulted mostly 
from an increase in infertile eggs and was not clearly related to a change 
in the inviable class. 

Since field-collected females were older and probably had laid some 
eggs before capture, it was more difficult to get samples of eggs for 
sufficiently long periods for analyses. Females from neighboring pop- 
ulations (northern Michigan, northern Wisconsin and so on) were grouped 
together without regard to year because there were few differences 
between years for lab females. The results for field females were similar 
to those for lab females. Females from both P. g. glaucus and P. g. 
canadensis populations showed significant declines in viability after 
three samples (Table 3). The decline was not significant for P. g. can- 
adensis females with four samples each (Table 4). This probably rep- 
resents random variability between the very low viabilities of the third 
and fourth samples (Table 2). The increases in infertility were significant 
for both groups for three samples and for P. g. glaucus for four samples. 
Again, there was no pattern in inviable eggs with sample. The signif- 
icantly nonhomogeneous result for P. g. canadensis females with four 
samples was a decrease in inviable eggs from sample to sample. 

The significant sequential decline in egg viability could be related 
to either the number of eggs that had been oviposited already or the 


VOLUME 41, NUMBER 2 87 


TABLE2. Average egg viability (percent hatching) for each of four sequential samples 
from P. glaucus females. Top values are means; lower values are SD. The first sample 
size (n,) is for the first three means; the second (n,) is for the fourth. 


Sample no. 
Group n, Ny ] 2 3 4 
1984 lab female-— 18 14 79.0 67.6 58.5 46.9 
lab male 23.4 iP 34.8 32.8 
1984 lab female-— 20 i, 74.2 56.9 45.4 40.2 
field male 29.6 26.1 32.1 31.8 
1985 lab female-— 18 13 91.2 81.0 64.1 46.4 
field male jel ker 29.1 35.6 3 ew 
P. g. glaucus DI 11 87.8 70.1 50.9 37.0 
field females Wan 26.8 39.6 41.1 
P. g. canadensis 21 1 79.3 38.9 28.7 38.2 
field females 24.3 Sie 33.0 ADs 


time interval since mating. To distinguish between the two possible 
causes, lab females were divided into those that laid fewer than 125 
eggs and those that laid more than 125 in the first two samples. For 
lab females mated to lab males, the viability of eggs in the first two 
samples combined did not differ significantly for the two groups (Mann- 
Whitney U, P > 0.50, Table 5). in addition, viability of third-sample 
eggs was independent of the number of eggs previously oviposited by 
the females (Mann-Whitney U, P > 0.50, Table 5). For lab females 
mated to field males, egg viability was actually higher for females that 
had laid more eggs. The differences were significant for the first two 
samples combined and for the third samples (Mann-Whitney U, both 
P’s < 0.05). Therefore, any decline in egg viability was not the result 
of females using up their supplies of sperm as they laid more eggs. The 
potential for a single insemination to fertilize a large number of eggs 
is illustrated by two females that laid 290 and 306 eggs within 5 days 


TABLE 8. Analyses for homogeneity for each category of eggs for three sequential 
samples from P. glaucus females. Mean interval is number of days between mating or 
capture and oviposition of the last sample. Probabilities are for a more extreme value of 
the Friedman two-way analysis of variance statistic. 








Interval (no. days) P 





Group n Mean SD Viable Inviable Infertile 
1984 lab female—lab male 18 6.9 1.6 0.05 N.S. n.s. 
1984 lab female—field male 20 Tet 2 0.01 0.05 0.05 
1985 lab female-field male 18 7.9 2.0 0.01 n.s. 0.01 
P. g. glaucus field females 21 10.0 2.6 0.02 N.S. 0.01 


P. g. canadensis field females 21 8.3 RE 0.001 n.s. 0.001 





88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Analyses for homogeneity for each category of eggs for four sequential 
samples from P. glaucus females. Mean interval is number of days between mating or 
capture and the oviposition of the last sample. Probabilities are for a more extreme value 
of the Friedman two-way analysis of variance statistic. 





Interval (no. days) P 
Group n Mean SD Viable Inviable Infertile 
1984 lab female—-lab male 14 10.3 2, if 0.001 N.S. 0.001 
1984 lab female—field male WZ Ted 3.4 0.01 N.S. 0.05 
1985 lab female-—field male 13 11.8 3.0 0.01 nS; 0.01 
P. g. glaucus field males il 12.5 2.8 0.01 ns. 0.001 
P. g. canadensis field females 12) 10.9 3.4 ns. 0.05 nS. 





after mating. The viabilities for these two clutches were 90.0% and 
97.1%, respectively. 

Lifetime oviposition was high for females in this study. Lab P. g. 
glaucus females laid an average of 181.2 eggs.+ 77.2 (n = 36). Yellow- 
and dark-morph individuals did not differ significantly in number of 
eggs laid. Field-collected P. g. glaucus laid 141.1 + 61.4 eggs (n = 21), 
and P. g. canadensis laid 130.0 + 44.0 eggs (n = 23). These values did 
not differ significantly (t-test, P > 0.20), but both were significantly 
less than the value for lab females (t-test, P < 0.05 and 0.01, respec- 
tively). 

Since the decline in viability was significant using the Friedman two- 
way analysis of variance, a multiple comparison analysis was applied 
to the ranked samples to determine which were significantly different 
from the others. Using a modified Tukey procedure (Zar 1984), a general 
pattern emerged. Females mated to lab males had fourth samples that 
were significantly lower in viability than the first two. In both groups 
of females hand-paired to field males, the first sample was significantly 


TABLE 5. Viability (percent hatching) related to number of eggs laid by P. glaucus. 
Females were grouped by oviposition during the first two sample intervals. Means for 
oviposition and viability are given for the first two samples combined and for viability 
for the third sample. 


Viability (percent hatching) 


No. of eggs ~ Twocombined  ~—=«*‘ThirdS 
Female group n Mean SD. 5 Mean 5) SD) 2 i McananODIO 
Lab female-lab male 
Less than 125 eggs 8 80.6 29.5 77.8 15.7 C2ZiGn wien 
More than 125 eggs MO eA 16 69.6 29.5 55.2 40.5 
Lab female-field male 
Less than 125 eggs 24 74.2 20.9 69.6 22.4 44.2 34.8 


More than 125 eggs 14 isto). oul 84.6 13.3 yale PAT 





VOLUME 41, NUMBER 2 89 


higher than the fourth sample. This suggests that the declines occurred 
sooner in females that had mated with males that may have mated 
before. For field-collected P. g. glaucus females, the first sample was 
significantly higher than the last two for both three- and four-sample 
analyses. For P. g. canadensis, the first sample was significantly higher 
than the last two for three samples per female. Since field-collected 
females had mated some time before capture, this reflects the longer 
intervals between mating and completion of oviposition for these fe- 
males (Tables 3, 4). 

Not all matings produced high levels of fertility and hatching. A 
proportion of the field-collected females that contained at least one 
spermatophore and laid 15 or more eggs laid no viable eggs. In 1985, 
21.4% of 28 Ohio P. g. glaucus females produced clutches with total 
hatching failure. In 1986, 21.3% of 89 Ohio females also produced no 
viable eggs. A similar sample of 28 P. g. canadensis females had a 
10.7% total failure rate. Comparable results are illustrated by 57 hand- 
pairings using males collected in Adams Co., Ohio. In 5 cases (8.8%), 
no spermatophore was transferred. An additional 12 females (21.0%) 
with spermatophores laid fewer than 5 eggs. Nine pairings (15.8%) 
resulted in entirely infertile eggs even though a spermatophore had 
been transferred. The remaining 31 pairings averaged 62.4% + 23.38% 
larval hatch. Thus, for hand-paired females that had a spermatophore 
and laid eggs, 22.5% produced clutches with total hatching failure. 

An additional mating often had a substantial effect on the viability 
of eggs laid subsequently. The effect of additional matings on hatching 
rate was investigated by comparing the first samples of eggs from field- 
collected Ohio females that mated once and that mated more than once. 
Females that had total hatching failure were excluded. Fresh singly- 
mated females had a mean hatching rate of 64.2% + 29.8% (n = 26) 
which was significantly higher than the 35.5% + 28.1% of 5 worn singly- 
mated females (Mann-Whitney U, P < 0.05). Worn multiply-mated 
females had a mean hatching rate of 69.4% + 27.0% (n = 14). This 
was not different from that of fresh singly-mated females but was 
significantly higher than worn singly-mated females (Mann-Whitney 
U, P > 0.30 and <0.05, respectively). 


DISCUSSION 


Analysis of spermatophore counts for the Ohio P. g. glaucus docu- 
ments the highest mean value for any population of this species studied 
to date. Not only is the mean nearly 2 spermatophores per female, but 
the proportion (65.4%) of the females that had mated more than once 
is also higher than any other reports for this species (Drummond 1984). 
Burns (1968) reported a mean of 1.75 spermatophores per female for 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


one population and a multiple-mating percent of 63% for another pop- 
ulation of P. g. glaucus. Number of spermatophores carried by Ohio 
females increased significantly with wing wear (age). This agrees with 
results of Platt et al. (1984) and helps to explain the only low mean 
‘spermatophore values reported for this species. Pliske (1972, 1978) 
reported means of about 1.15 spermatophores per female, but his sam- 
ples of females were considerably less worn. 

As Levin (1973) demonstrated, a single normal insemination is suf- 
ficient to fertilize all the eggs a P. glaucus female is likely to lay. Our 
study confirms this even for females that oviposited over 50% more 
eggs on average than those in Levin’s study as long as they were laid 
within a week of mating. In addition, our experiments document the 
potential for the sperm of one mating to produce fertilities of over 95% 
for females laying 300 or more eggs. That one normal insemination is 
sufficient also generally agrees with reports for Papilio zelicaon Lucas 
(Sims 1979), P. polyxenes (Lederhouse 1981), and Pieris rapae crucivora 
Bsd. (Suzuki 1979). 

In contrast to Levin (1973), however, we document two situations in 
which an additional mating would be selectively advantageous for a 
Papilio glaucus female. First, a particular mating by a female is not 
always effective in fertilizing eggs. Second, even for a highly effective 
mating, the viability of the eggs of singly-mated females declines con- 
siderably with time regardless of oviposition rate. 

In P. zelicaon, a newly emerged male or one that has mated fre- 
quently produces an abnormally small spermatophore often with a low 
sperm count (Sims 1979). In our study, 22.5% of hand-paired females 
that had spermatophores and laid eggs had total hatching failure. Another 
15% of hand-pairings with some eggs hatching resulted in viabilities of 
less than 50% for the first two samples of eggs. Between 10% and 20% 
of field-collected females carrying at least one spermatophore laid no 
viable eggs. Although the complete hatching failure of some field- 
collected females may have resulted from a decline in viability with 
the time interval between mating and oviposition in the laboratory, our 
results suggest that the failure rate for natural matings is considerable 
(Drummond 1984). 

The potential for long-term sperm storage is present in butterflies. 
Females of several species of Heliconius are capable of laying fertile 
eggs over 4-6 month lifespans (Dunlap-Pianka et al. 1977) although 
they usually mated only once (Ehrlich & Ehrlich 1978). However, in 
relatively short-lived species such as various swallowtails the sperm 
supply does deteriorate fairly quickly. Remington (1959) noted that 
later samples of eggs from hand-paired Papilio females had low fertility. 
In P. polyxenes, this decline was time-related but independent of the 


VOLUME 41, NUMBER 2 9] 


number of eggs that had been laid (Lederhouse 1981). In Euphydryas 
editha Bsd., a single mating was effective for only 7 to 10 days (Labine 
1966). 

Egg viabilities and fertilities of singly-mated tiger swallowtails de- 
creased over time. Our results for P. glaucus appear at odds with those 
of Levin (1973). However, this is likely the result of the average lifespan 
of 5.4 days for the females in his study. There was a decline in fertility 
for this period in Levin’s study, but it was not significant. Both male 
and female P. glaucus live up to two weeks in the field (Berger 1986, 
Lederhouse 1982, unpubl. data). The 7- to 13-day average lifespans of 
the females in our study are more representative of a proportion of the 
females in the field. Females of short-lived butterfly species in thermally 
limiting environments may not benefit from mating more than once 
(Wiklund 1977a, 1982). 

Egg viability decreased more gradually throughout the first week for 
females mated to virgin males. Only the fourth sample was significantly 
lower than the first for these females. These values declined sooner and 
more if the males were field-collected. This suggests that these males 
provided smaller spermatophores, possibly as a result of depletion of 
fluids from a previous mating, as is the case for multiply-mating P. 
zelicaon males (Sims 1979). Since spermatophores much smaller than 
average do not produce a mating refractory period (Sugawara 1979), 
females in the field could correct deficiencies by mating an additional 
time. Indeed, 50% of the Ohio females in the two freshest wear classes 
that had mated more than once had a first spermatophore that was 
considerably smaller than average (R. Lederhouse unpubl. data). 

Declines in egg viability were substantial for all groups of females 
amounting to a reduction to half the initial viability level by 10 days 
after pairing. Although an additional mating does not guarantee higher 
viability, multiply-mated worn females had significantly higher via- 
bilities of their eggs than singly-mated worn individuals. The average 
viability values for multiply-mated worn females did not differ from 
those of singly-mated fresh females. For females that live more than a 
week, an additional mating could be very advantageous since it is likely 
to restore viability levels to average values. Females of Pieris brassicae 
L. and P. rapae crucivora typically mate a second time a week after 
the first mating (David & Gardiner 1961, Suzuki 1979). 

The rate of mating failure in natural copulations (10-20%) and the 
decline over time in the viability of eggs fertilized by a single male 
combine to make the probability of remating important in the tiger 
swallowtail. Sexual selection by males for yellow females could provide 
a differential favoring this morph. The dark morph females should on 
average live longer because of their mimetic advantage. However, if 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the additional eggs they might lay during this period are largely infertile 
resulting from lower or slower remating rates, the advantage of their 
greater longevity would be reduced. The importance of sexual selection 
in maintaining the color polymorphism in the tiger swallowtail will not 
be resolved using spermatophore count data alone because of the lim- 
itations of this technique. However, mating frequency clearly affects 
lifetime reproductive success in P. glaucus. Suggestions that female 
butterflies should mate only once (Wiklund 1977b) must be limited to 
short-lived species. In longer-lived species, selection should favor regular 
mating intervals or more effective sperm storage. Such factors warrant 
further investigation in long-lived species. 


ACKNOWLEDGMENTS 


We thank S. G. Codella, B. A. Drummond, R. H. Hagen, D. D. Murphy, and S. J. 
Rockey for helpful comments on drafts of this paper. We are grateful to M. Evans, Y. 
Allen, J. Johnson, C. Plzak, E. Schuh, J. Sibenhorn, J. Thorne, V. Viegut, and D. Ware 
for their assistance. Support for this research was provided in part by NSF grants DEB 
79-21749 and BSR 8306060; USDA grant 85CRCR-1-1598 and the Graduate School and 
the College of Agriculture and Life Sciences (Hatch 5134) of the University of Wisconsin, 
Madison. 


LITERATURE CITED 


BERGER, T. A. 1986. Habitat use and reproductive ecology of the eastern tiger swal- 
lowtail, Papilio glaucus L. Dissertation, Rutgers University, Newark, New Jersey, 
USA. 109 pp. Diss. Abstr. Int. microfilm order No. 8616563. 

BROWER, J. VZ. 1958. Experimental studies of mimicry in some North American but- 
terflies. Part IJ. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus. 
Evolution 12:123-136. 

BROWER, L. P. & J. VZ. BROWER. 1962. The relative abundance of model and mimic 
butterflies in natural populations of the Battus philenor mimicry complex. Ecology 
43:154-158. 

BURNS, J. M. 1966. Preferential mating versus mimicry: Disruptive selection and sex- 
limited dimorphism in Papilio glaucus. Science 153:551-553. 

1967. Reply to: Selective forces in Papilio glaucus. Science 156:534. 

1968. Mating frequency in natural populations of skippers and butterflies as 
determined by spermatophore counts. Proc. Nat. Acad. Sci. 61:852-859. 

CLARKE, C. A. & P. M. SHEPPARD. 1962. The genetics of the mimetic butterfly Papilio 
glaucus. Ecology 43:159-161. 

Davin, W. A. L. & B. O. C. GARDINER. 1961. The mating behaviour of Pieris brassicae 
(L.) in a laboratory culture. Bull. Entomol. Res. 52:263-280. 

DRUMMOND, B. A. 1984. Multiple mating and sperm competition in the Lepidoptera, 
pp. 291-370. In Smith, R. L. (ed.), Sperm competition and the evolution of animal 
mating systems. Academic Press. 

DUNLAP-PIANKA, H., C. L. Boccs & L. E. GILBERT. 1977. Ovarian dynamics in heli- 
coniine butterflies: Programmed senescence versus eternal youth. Science 197:487— 
490. 

EHRLICH, A. H. & P. R. EHRLICH. 1978. Reproductive strategies in the butterflies: I. 
Mating frequency, plugging, and egg number. J. Kansas Entomol. Soc. 51:666—697. 

LABINE, P. A. 1966. The population biology of the butterfly, Euphydryas editha. IV. 
Sperm precedence—A preliminary report. Evolution 20:580-586. 

LEDERHOUSE, R. C. 1978. Territorial behavior and reproductive ecology of the black 








VOLUME 41, NUMBER 2 93 


swallowtail butterfly, Papilio polyxenes asterius Stoll. Dissertation, Cornell Univer- 

sity, Ithaca, New York. 155 pp. Diss. Abstr. Int. order No. 7902342. 

1981. The effect of female mating frequency on egg fertility in the black 

swallowtail, Papilio polyxenes asterius (Papilionidae). J. Lepid. Soc. 35:266-277. 

1982. Factors affecting equal catchability in two swallowtail butterflies, Papilio 
polyxenes and P. glaucus. Ecol. Entomol. 7:379-383. 

LEDERHOUSE, R. C. & J. M. ScrIBER. 1987. Increased relative frequency of dark morph 
females in the tiger swallowtail, Papilio glaucus (Lepidoptera: Papilionidae) in S-cen- 
tral Florida. Am. Mid]. Nat. in press. 

LEVIN, M. P. 1978. Preferential mating and the maintenance of the sex-limited di- 
morphism in Papilio glaucus: Evidence from laboratory matings. Evolution 27:257- 
264. 

MAKIELSKI, S. K. 1972. Polymorphism in Papilio glaucus L. (Papilionidae): Maintenance 
of the female ancestral form. J. Lepid. Sco. 26:109-111. 

PLATT, A. P., S. J. HARRISON & T. F. WILLIAMS. 1984. Absence of differential mate 
selection in the North American tiger swallowtail, Papilio glaucus, pp. 245-250. In 
Vane-Wright, R. I. and P. R. Ackery (eds.), The biology of butterflies. Sympos. 11. 
Academic Press, London. 

PLISKE, T. E. 1972. Sexual selection and dimorphism in female tiger swallowtails, Papilio 
glaucus L. (Lepidoptera: Papilionidae): A reappraisal. Ann. Entomol. Soc. Am. 65: 
1267-1270. 

1973. Factors determining mating frequencies in some New World butterflies 
and skippers. Ann. Entomol. Soc. Am. 66:164-169. 

PRouT, T. 1967. Selective forces in Papilio glaucus. Science 156:534. 

REMINGTON, C. L. 1959. Wide experimental crosses between Papilio xuthus and other 
species. J. Lepid. Soc. 13:151-164. 

SCRIBER, J. M. 1985. The ecological and genetic factors determining geographic limits 
to the dark morph polymorphism in Papilio glaucus. Bull. Ecol. Soc. Am. 66:267. 

SIEGEL, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-Hill, 
New York. 312 pp. 

Sims, S. R. 1979. Aspects of mating frequency and reproductive maturity in Papilio 
zelicaon. Am. Midl. Nat. 102:36-50. 

SUGAWARA, T. 1979. Stretch reception in the bursa copulatrix of the butterfly, Pieris 
rapae crucivora, and its role in behaviour. J. Comp. Physiol. A. 130:191-199. 

SUZUKI, Y. 1979. Mating frequency in females of the small cabbage white, Pieris rapae 
crucivora Boisduval (Lepidoptera: Pieridae). Kontyt 47:335-339. 

WIKLUND, C. 1977a. Oviposition, feeding, and spatial separation of breeding and for- 
aging habitats in a population of Leptidea sinapis (Lepidoptera). Oikos 28:56-68. 

1977b. Courtship behaviour in relation to female monogamy in Leptidea sinapis 

(Lepidoptera). Oikos 29:275-283. 

1982. Behavioural shift from courtship solicitation to mate avoidance in female 
ringlet butterflies (Aphantopus hyperanthus) after copulation. Anim. Behv. 30:790- 
793. 

ZAR, J. H. 1984. Biostatistical analysis. 2nd ed. Prentice-Hall, Englewood Cliffs, New 
Jersey. 718 pp. 

















Received for publication 15 August 1986; accepted 14 January 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 94-97 


DEMOGRAPHY OF THE UNSILVERED MORPH OF 
SPEYERIA MORMONIA IN COLORADO 


CAROL L. BOGGS 


Rocky Mountain Biological Laboratory, Box 519, 
Crested Butte, Colorado 81224 and 
Department of Biological Sciences, Stanford University, 
Stanford, California 94305 


ABSTRACT. The demographies of unsilvered and silvered morphs of Speyeria mor- 
monia do not differ greatly in a Colorado population with a very low frequency of 
unsilvered animals. The frequency of the unsilvered morph is on the order of that expected 
if the unsilvered morph were due to a recessive allele with fitness zero, which is maintained 
only by mutation. The lack of detectable difference between morphs in demographic 
parameters suggests, however, that other factors control the frequency of the polymor- 
phism. 


Additional key words: Nymphalidae, polymorphism. 


The silver spots on the underside of the wings of Speyeria mormonia 
Edwards (Nymphalidae) consist of modified scales. These silver spots 
are absent in some individuals, being replaced by buff colored spots. 
The percentage of unsilvered morphs in a population varies substantially 
among populations (Remington 1956). 

It is not known whether the silvered-unsilvered difference is actually 
heritable, although Remington (1956) speculates that it may be con- 
trolled by one autosomal locus. The frequency of the unsilvered morph 
in a population may therefore be determined by any of a number of 
factors, including mutation rate, natural selection, drift, and—if the 
dimorphism should prove to be due to environmental plasticity—en- 
vironmental characteristics. 

Even in the absence of genetic information, we can begin to under- 
stand something of the forces governing the frequency of the morphs 
in a population by comparing demographic parameters for the two 
morphs. This is particularly true for populations in which the frequency 
of one morph is extremely low. In such cases, an extreme discordance 
in demographic parameters for the two morphs would suggest that 
extreme selection pressures relating to the unsilvered and silvered phe- 
notypes exist in the adult stage. In the particular population examined 
here, the frequency of the unsilvered morph is in the range of that 
expected if unsilvered is due to a recessive allele whose fitness is zero 
and whose frequency in the population is thus determined strictly by 
the mutation rate. 

Here I compare demographic data obtained over four years for 
unsilvered and silvered morphs in a Colorado population of S. mor- 
monia. Recapture rates and maximum longevities (indices of survival), 
dispersal, and sex ratio are compared for the two morphs. I also examine 


VOLUME 41, NUMBER 2 95 


yearly shifts in percentage of unsilvered morphs in the population. 
While the sample size of unsilvered animals is by necessity quite small 
in a population with such a low frequency of unsilvered morphs, dif- 
ferences in values for demographic parameters are expected to be 
dramatically large if mutation balanced by low adult survival yielding 
a fitness of zero is responsible for the unsilvered morph’s low frequency. 
The results thus give a first indication of the type and extent of differ- 
ences between the morphs. 


MATERIALS AND METHODS 


The study was conducted near Gothic, Gunnison Co., Colorado. The 
study area consists of fescue grassland (Langenheim 1962), sometimes 
bordered by stands of aspen or spruce. Elevation varies from 2880 to 
2970 m. Most lower slopes face SW, while upper slopes face NW. The 
study area was divided into topographic sites for calculation of dispersal 
distances. Total area sampled in 1979 was 6.0 ha; in 1980, 24.7 ha; in 
1981, 5.6 ha; and in 1982, 6.3 ha (see Boggs 1987 for further detail). 

Standard mark-release-recapture techniques were used in this study 
each summer between 1979 and 1982. Date and time of capture, but- 
terfly number, sex and site of capture were recorded. Captured indi- 
viduals were immediately released at the site of capture. Recaptures 
of individuals on the same day in the same site were not recorded. 
Dispersal distances were measured between centers of sites. 


RESULTS AND DISCUSSION 


Frequency of unsilvered animals captured in this population varied 
between 0% and 0.2% (Table 1), and did not differ significantly among 
years (Table 2). In 1981, the one unsilvered animal caught had half of 
each normally silvered spot unsilvered. 

If the frequency of unsilvered animals in the population were con- 
trolled by the mutation rate alone, with the fitness of homozygous 
recessive unsilvered animals equal to zero, then the square of the fre- 
quency of unsilvered would equal the mutation rate. Mutation rates 
for this population would therefore be 5.1 x 10~°in 1980, 1.4 x 107° 
in 1981 and 4.7 x 10~° in 1982, which are in the range reported for 
whole-locus mutation rates in Drosophila (Crow 1986). 

Given a total density for this population of about 2000/ha, and an 
average dispersal distance of about 175 m (Boggs 1987), the lack of 
significant difference among years in frequency does not rule out ge- 
netic drift as a factor affecting frequencies of the morphs. For example, 
the time to fixation for a neutral mutant is approximately four times 
the effective population size (Crow 1986); the change in frequency in 
four generations under these circumstances would not necessarily be 
large enough to detect without a much larger sample size. 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Number and sex ratio of silvered and unsilvered S. mormonia marked per 
year. 


Sex ratio 
Total no. Total no. 

Year silvered unsilvered Silvered Unsilvered 
1979 090 0 Sy Well — 
1980 2646 6 oor et 
1981 846 Le Wall 1:0 
1982 2762 6 2.4:1 oul 

Total 6844 13 3.2:1 2.2:1 


* Half-silvered. 


The sex ratio of unsilvered animals showed no consistent pattern 
(Table 1). Taken over all four years, the sex ratio of unsilvered animals 
did not differ significantly from that of silvered animals (Table 2). This 
is consistent with the hypothesis of determination of silvering by an 
autosomal locus. 

Unsilvered individuals showed no pronounced difference from the 
silvered in other demographic attributes. Based on recapture frequen- 
cies for silvered animals (Boggs 1987), I expected to recapture 0.1, 0, 
and 0.2 unsilvered females in 1980, 1981, and 1982, respectively. None 
were ever recaptured. I expected to recapture 0.4, 0.1, and 1.8 unsil- 
vered males in 1980, 1981, and 1982, respectively; 1, 0, and 2 males 
were recaptured in 1980, 1981, and 1982, respectively. The extremely 
close correspondence of these data to expectations over several years, 
in spite of the small numbers involved, indicates that survival and 
catchability of unsilvered individuals paralleled that of silvered but- 
terflies among years. Ordinary statistical tests are precluded by the 
small numbers intrinsic to the situation. However, it is clear that, com- 
pared to silvered morphs, the unsilvered did not suffer massively greater 
mortality in the adult stage. 

Maximum residence time is indicated by the interval between first 
and last capture. Maximum residence time in 1980 was 8 days for the 
one unsilvered male recaptured, whereas it was 15 and 28 days for the 
two unsilvered males recaptured in 1982. These values are not incon- 
sistent with values for silvered individuals. For such butterflies, daily 
survival rates were lower in 1980 than in 1982, and maximum residence 
seen was 28 and 40 days, respectively, in 1980 and 1982 (Boggs 1987). 

Dispersal characteristics of unsilvered and silvered morphs did not 
differ greatly either. The recaptured male in 1980 was recaptured in 
the same site as originally caught, yielding a dispersal distance of 0 m. 
Distances moved by the recaptured males in 1982 were 180 m and 580 
m. Average distance moved by silvered morphs was about 175 m, with 
60-80% of the recaptured animals dispersing (Boggs 1987). 


VOLUME 41, NUMBER 2 O7 


TABLE 2. Tests of significance of differences among years in percentage of unsilvered 
morphs in the population, and of differences between morphs in the four year total sex 
ratio. The test statistic, x*, Goldstein’s (1964) exact binomial test for differences of pro- 
portions, equals the difference between the proportions divided by a pooled standard 
deviation, and is compared to values for t.,. 95% confidence intervals for the difference 
between proportions indicate the amount of difference which could exist but not be 
detected as significant at P = 0.05. 


95% confidence interval for 
Test xe P difference between percentages 


Percentage unsilvered between years 


1979 (0%) vs. 1980 (0.23%) = IIe: >0.05 —0.15% to 0.61% 
1979 (0%) vs. 1981 (0.12%) —0.835 >0.05 —0.16% to 0.39% 
1979 (0%) vs. 1982 (0.22%) leloe >0.05 —0.16% to 0.59% 
1980 (0.23%) vs. 1981 (0.12%) 0.614 >0.05 —0.24% to 0.45% 
1980 (0.23%) vs. 1982 (0.22%) 0.074 >0.05 —0.24% to 0.26% 
1981 (0.12%) vs. 1982 (0.22%) —0.573 >0.05 —0.24% to 0.44% 


Percentage males between morphs 


Silvered (76.3%) vs. unsilvered 
(69.2%) 0.595 >0.05 —]16.2% to 30.2% 


There was thus no evidence for an extreme survival-residence dis- 
advantage of the unsilvered morph. This is contrary to expectations if 
the frequency of the unsilvered morph is maintained by mutation in 
combination with an extreme selective disadvantage. However, the 
evidence presented here does not completely rule out maintenance of 
the unsilvered morph by a balance between mutation and selection, as, 
for example, ability to successfully lay eggs or mate was not examined. 


ACKNOWLEDGMENTS 


This work was financially supported by the Bache Fund of the National Academy of 
Sciences, the American Philosophical Society, the Roosevelt Memorial Fund, A. and L. 
McCarley, the Mobil Foundation, R. and L. Watt, and J. and R. A. Boggs. E. Maxwell, 
J. Delgado, L. Nordby, R. Loveland, D. Price, S. Sonnad and the 1982 Earthwatch Team 
“Rocky Mountain High” assisted in the mark-release-recapture study. B. Bench and the 
Enders family kindly gave permission to conduct part of the study on their land. This 
work benefitted from discussion with C. Remington and especially W. Watt. 


LITERATURE CITED 


Boccs, C. 1987. Within population variation in the demography of Speyeria mormonia 
(Lepidoptera: Nymphalidae). Holarctic Ecology. In press. 

Crow, J. 1986. Basic concepts in population, quantitative and evolutionary genetics. 
W. H. Freeman, New York. 278 pp. 

GOLDSTEIN, A. 1964. Biostatistics. MacMillan, New York. 272 pp. 

LANGENHEIM, J. 1962. Vegetation and environmental patterns in the Crested Butte 
area, Gunnison County, Colorado. Ecol. Monogr. 32:249-285. 

REMINGTON, C. L. 1956. Genetics of populations of Lepidoptera. Proc. 10th Int. Congress 
Entomol. 2:787—-828. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 98-103 


XANTHORHOE CLARKEATA (GEOMETRIDAE), 
A NEW SPECIES AND POSSIBLE ENDEMIC OF THE 
QUEEN CHARLOTTE ISLANDS, BRITISH COLUMBIA 


DOUGLAS C. FERGUSON 


Systematic Entomology Laboratory, Agricultural Research Service, USDA, 
% U.S. National Museum of Natural History, Washington, D.C. 20560 


ABSTRACT. Xanthorhoe clarkeata, new species, of which 159 specimens were col- 
lected in 2 days in the Alpine Zone on Graham Island, is the first species of Lepidoptera 
reported only from the Queen Charlotte Islands. It is related to a holarctic species, X. 
abrasaria, but is easily distinguished by its usually reddish-brown coloring, greater color 
variability, strongly pectinate male antennae, and diurnal flight, as well as by genitalic 
differences. The new species may be a Pleistocene refugial relict. This aspect is discussed 
relative to what has been published on other endemic animals and plants of these islands, 
and on the biological evidence for a refugium there during the last glaciation. 


Additional key words: taxonomy, biogeography, relict populations. 


On a visit to the Queen Charlotte Islands, British Columbia, in July 
1985, J. F. Gates Clarke of the U.S. National Museum of Natural History 
chartered a helicopter for two flights to the otherwise inaccessible moun- 
tains of the interior. His purpose was to investigate the Lepidoptera of 
the alpine tundra in these isolated ranges where no previous collections 
are known to have been made. One flight took him to 3100 ft (946 m) 
on a ridge near Mt. Brown, Graham Island, where he camped overnight. 
The other flight was a 1-day trip to a 3000 ft (915 m) ridge on Moresby 
Island. No butterflies and relatively few moths were present, but at the 
Mt. Brown locality a day-flying moth occurred in abundance. Dr. Clarke 
recognized it as something unusual, and he netted and pinned 159 
specimens. Although virtually swarming above tree line on Graham 
Island, this species was not seen in similar habitats and under similarly 
favorable weather conditions on Moresby Island. 

Comparison of these moths with all described North American species 
and subspecies and most palearctic species of Xanthorhoe, as well as 
with superficially similar taxa in other genera, leaves little doubt that 
it is undescribed. It is not a cryptic species but a distinctive one. I am 
pleased to name it after Jack Clarke, whose dedicated investigation of 
the Lepidoptera of the Queen Charlotte Islands led to its discovery and 
who collected all known specimens. Illustrations are by the author. 


Xanthorhoe clarkeata Ferguson, new species 


(Figs. 1-11) 


Diagnosis. A variable day-flying species found above tree line on the mountains of the 
Queen Charlotte Islands, British Columbia. The most characteristic feature of the forewing 
pattern is the banding of the median space, in which the antemedial line is followed and 
the postmedial line preceded by fairly wide and regular light reddish brown to nearly 


VOLUME 41, NUMBER 2 99 





SS 


Fic. 1. Male genitalia of Xanthorhoe clarkeata. Aedeagus removed and shown in 
ventral view. 


black bands that tend to constrict the much paler grayish central space between them. 
A small dark discal spot usually marks the center of what remains of the pale median 
space. Genitalia most resemble those of X. abrasaria (Herrich-Schaeffer) in general con- 
figuration, differing mainly in the shape or proportions of components, although one 
would hardly guess from external appearances that the two species were closely related. 
The usually reddish brown clarkeata has narrower, more pointed wings than the pre- 
dominantly gray and black abrasaria, and its male antennae are bipectinate (heavily 
setose but not bipectinate in abrasaria). Xanthorhoe clarkeata is about the size of north- 
eastern American abrasaria (subspecies congregata Walker). The forewing pattern, as 
well as the pectinate antennae, at first led me to compare clarkeata with the European 
Scotopteryx peribolata Hubner, but the genitalia are in no way similar. 

Further description. Male antenna bipectinate, with black branches; longest branches 
about equal in length to 2% antennal segments; ventral branches twice length of dorsal 
ones; branches conspicuously setose, shaft covered with brown scales to tip; female antenna 
simple, slender. Palpus rough scaled, light brown, exceeding front for about half its length, 
somewhat decumbent toward end. Front slightly convex, rough scaled, matching color 
of palpi; gena unscaled. Eye reduced to % width of front in male, % width of front in 
female (width of eye equal to or greater than width of front in both sexes of abrasaria); 
eye finely and sparsely setose in both sexes. Chaetosemata large but not meeting mesially. 
Tongue well developed, dark brown, nearly black. Legs normal, fairly slender, with two 
pairs of hindtibial spurs in both sexes. Vestiture of legs and body a nearly uniform mixture 
of pale and darker gray-brown scales. 

Wings appearing narrower than those of abrasaria because forewing is apically more 


100 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





Fic. 2. Female genitalia of Xanthorhoe clarkeata. 


produced, that of female ending in slightly acute apex. Ground color of forewing pale 
bluish gray, but in nearly all males and some females this shade mostly obscured by 
darker gray-brown to bright reddish brown suffusion; pale basal and antemedial lines 
roughly parallel, convex, angled or bending inward toward costa, usually enclosing a 
grayish to rust-colored band contrastingly paler than basal band or proximal dark band 
in median space beyond; median space with its light-gray central area variably constricted 
by dark-brown to reddish-brown transverse bands that closely follow antemedial and 
postmedial lines; pale central area may remain a continuous medial gray band from costa 
to inner margin (Figs. 3, 9), be pinched off to form separate gray areas near costa and 
inner margin (Fig. 4) or one discal patch bearing the dark discal spot in middle (Fig. 7); 
postmedial line whitish, regular or slightly irregular but not as sinuous as that of X. 
abrasaria, convex near middle; dusky to rust-brown distal third of forewing often almost 
evenly divided by crenulate or dentate gray subterminal line parallel to outer margin; 
terminal line a series of double dark dots, one on each side of each vein; apex marked 
by small, oblique blackish dash directed toward or just behind apex. Hindwing darkened 


VOLUME 41, NUMBER 2 101 





Fics. 8-11. Xanthorhoe clarkeata. 3, Holotype; 4, 5, Male paratypes, 28 July 1985; 
6, 7, Male paratypes, 27 July 1985; 8, Male paratype, 28 July 1985; 9-11, Female 
paratypes, 27 July 1985. 


by dusky suffusion, marked by faint discal spot, by diffuse pale postmedial and subterminal 
bands paralleling outer margin, and series of terminal dots like those of forewing. Fringes 
ot both wings dusky to light gray or pale yellowish brown. Females usually much paler 
and more extensively grayish than males, although some are predominantly brown, and 
they usually have the light-gray median band of forewing unconstricted and much wider. 
Undersides of both wings dusky to reddish brown, with diffuse, darker postmedial bands 
with paler shading beyond, and often a diffuse subterminal, also pale-shaded outwardly. 
Length of forewing: holotype, 12.0 mm, other males, 11.0-14.0 mm (N = 144); females, 
11.5-138.0 mm (N = 12). 

Male genitalia (Fig. 1) (N = 2). Closest to those of X. abrasaria but with several 
conspicuous and numerous lesser differences; most notably: costa of valve with one or 
two dentate, subapical processes on inner side; membranous lobe of valve less ample, 
more flattened; costa of valve stouter, more heavily developed; large papillate process of 
juxta erect, recurved, not recumbent; saccus knoblike, rounded, not acuminate and point- 
ed; manica more heavily spined than that of abrasaria; and vesica with similar two groups 
of cornuti, but distal ones longer than proximal ones, the reverse of abrasaria. 

Female genitalia (Fig. 2) (N = 1). Differ from those of abrasaria in having larger, fully 
sclerotized ductus bursae about as long as bursa copulatrix, and with straight, transverse 
rim at ostium; a small signum only ™% the size of that in abrasaria; better developed 
anterior apophyses 2 or 3 times as long as those of abrasaria; with integument of segments 
7 and 8 dark pigmented, a difference often affecting the entire exoskeleton of diurnal 
moths. 

Types. Holotype 6, Ridge W of Mt. Brown, 3100’ [946 m], Graham Island, Queen 
Charlotte Islands, British Columbia, 27 July 1985, J. F. G. Clarke. Paratypes: 106 é, 12 
2, same data; 38 6, 1 2, same locality and collector but taken 28 July 1985. Holotype and 
most paratypes in U.S. National Museum of Natural History; some paratypes distributed 
to American Museum of Natural History, Canadian National Collection, British Museum 
(Natural History), Los Angeles County Museum of Natural History, Queen Charlotte 
Islands Museum at Skidegate Mission, Graham Island, and other collections. 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Remarks. In the description I compared Xanthorhoe clarkeata with X. abrasaria, a 
widespread, holarctic species with some obvious similarities in structure and wing pattern. 
The two are so easily distinguished by superficial features that I did not think it necessary 
to include figures of abrasaria. Illustrations of it may be found in Ferguson (1954:pl. 15, 
fig. 7), Morris (1980:pl. 29, fig. 10), and in many European works. An outline drawing 
of the male valve of X. abrasaria was given by Forbes (1948:148, fig. 158). 


BIOGEOGRAPHY 


Xanthorhoe clarkeata is the first species of Lepidoptera to be recorded 
only from the Queen Charlotte Islands. Such a restricted distribution 
is unusual because most alpine and subalpine Lepidoptera in northern 
North America are more widespread. Many are holarctic. Although 
this species might still prove to occur elsewhere, it seems equally likely 
to be a local endemic or relict. 

A few taxa endemic to the Queen Charlotte Islands have been rec- 
ognized in other groups such as Coleoptera (Lindroth 1962:67), am- 
phipod crustaceans (Bousfield 1958:64), and plants (Calder & Taylor 
1968:102, 103, Schofield 1969:174). Endemism in the plants is regarded 
as surprisingly high, with four endemic species and seven well-differ- 
entiated subspecies, and all but one are limited to about the same alpine 
habitat as X. clarkeata. Although geological evidence has not been 
reassuring, biologists have long maintained that a Pleistocene refugium 
must have existed in the Queen Charlotte Islands (Calder & Taylor 
1968:113, Schofield 1969, Randhawa & Beamish 1972). The concept 
that such a refugium persisted throughout the last (late Wisconsin) 
glaciation received new impetus from recent radiocarbon dating and 
analysis of plant remains in sediments and peat deposits by Warner et 
al. (1982), who showed that a diverse flora did exist near sea level on 
the E side of Graham Island 15,000 to 16,000 years ago and continuously 
thereafter. That was around the climax of the last glaciation, when the 
British Columbia mainland coast is believed to have been fully covered 
by glacial ice. The plant fossil deposits revealed the presence in the 
late Pleistocene of many of the same species that grow on the islands 
today, but some of them now only at higher elevations. 

Considering the growing biological evidence for a refugium in the 
Queen Charlotte Islands, a few endemic species or subspecies of Lep- 
idoptera were to be expected in addition to the previously known 
lycaenid, Lycaena mariposa charlottensis (Holland). Such an abundant 
and mobile species as X. clarkeata also might have spread back to the 
mainland in post-Pleistocene times, and I urge collectors to look for it 
elsewhere in the mountains of the N Pacific coast. The apparently 
endemic carabid beetle, Nebria charlottae (Lindroth 1962), has a close 
sister species in the Aleutians. Nothing comparably similar to clarkeata 
has yet appeared among the described and undescribed species of 


VOLUME 41, NUMBER 2 103 


Xanthorhoe from Alaska or the Aleutians, but those vast regions could 
hardly be described as well collected. 


LITERATURE CITED 


BOUSFIELD, E. L. 1958. Fresh-water amphipod crustaceans of glaciated North America. 
Can. Field Nat. 72:55-113. 

CALDER, J. A. & R. L. Taytor. 1968. Flora of the Queen Charlotte Islands. Pt. 1. 
Canada Dept. Agric. Res. Branch Monogr. 4. 659 pp. 

FERGUSON, D. C. 1954. The Lepidoptera of Nova Scotia. 1. Macrolepidoptera. Proc. 
Nova Scotian Inst. Sci. 23:161-375. 

ForBES, W. T. M. 1948. Lepidoptera of New York and neighboring states. Pt. 2. Cornell 
Univ. Agr. Exp. Sta. Mem. 274. 263 pp. 

LINDROTH, C. H. [1962]. The ground-beetles (Carabidae, excl. Cicindelidae) of Canada 
and Alaska. Pt. 2. Opusc. Entomol. Suppl. 20. 200 pp. 

Morris, R. F. 1980. Butterflies and moths of Newfoundland and Labradory. The Macro- 
lepidoptera. Canada Dept. Agric. Res. Branch Publ. 1691. 407 pp. 
RANDHAWA, A. S. & K. J. BEAMISH. 1972. The distribution of Saxifraga ferruginea and 
the problem of refugia in northwestern North America. Can. J. Bot. 50:79-87. 
SCHOFIELD, W. B. 1969. Phytogeography of northwestern North America: Bryophytes 
and vascular plants. Madrono 20:155-207. 

WARNER, B. G., R. W. MATHEWES & J. L. CLAGUE. 1982. Ice-free conditions on the 
Queen Charlotte Islands, British Columbia, at the height of the late Wisconsin gla- 
ciation. Science 218:675-677. 


Received for publication 6 February 1986; accepted 27 January 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 104-107 


A NEW SPECIES OF NEARCTIC BOMOLOCHA (NOCTUIDAE) 
FROM THE APPALACHIAN AREA 


LINDA BUTLER 


Division of Plant & Soil Sciences, P.O. Box 6108, West Virginia University, 
Morgantown, West Virginia 26506-6108 


ABSTRACT. Bomolocha appalachiensis, new species, is described from West Vir- 
ginia, Kentucky and North and South Carolina. The type series consists of four males 
and seven females. The species is easily distinguished from known Bomolocha by forewing 
color pattern and male genitalia. 


Additional key word: taxonomy. 


Five specimens of a unique Bomolocha were taken in three West 
Virginia counties between 1977 and 1979. The species appears to be 
undescribed, and an additional six specimens were located in collections. 
Nothing is known of its life history. | 


Bomolocha appalachiensis, new species 


(Figs. 1-7) 


Description. Male. Palpi twice length of head, strongly curving upward to horizontal 
above antennal bases; laterally compressed, heavily clothed with flat scales dorsally and 
laterally, scales tan, tipped with dark brown; palps clothed ventrally with flattened hairs, 
dark brown, flecked with tan; third palpal segment small and white tipped. Antennae 
filiform, with white scaling at pedicel and dorsally on all segments. Vestiture of head 
dark brown with some light brown scales; frontal tuft strong and conical. 

Vestiture of thorax and abdomen consisting of chocolate brown hairs and long flattened 
hairs mixed with elongated scales, concolorous with median area of wings. Basal tuft of 
thorax extending upward and strongly truncate. Abdominal tufts on segments 2-5, best 
developed on segments 3 and 4. Antemedian line of wings dark but not strongly con- 
trasting, sinuous and slightly excurved at top of cell and in fold (Fig. 1). Postmedian line 
sharply defined, margined apically with pale scales, with smoothly rounded bulges at cell 
and at fold, the latter bulge stronger. Median area of wing darker, richer brown than 
basal area. A series of dots between veins represents subterminal line; most specimens 
with obscure subterminal dots. Some specimens with faint row of terminal dots between 
veins. Wing beyond postmedian line sandy to grayish brown, darker apically; paler 
costoapical triangle above. Hindwings medium brown or grayish brown with more luteus 
fringe and dark brown terminal line. Forewing length: 15.5-16.5 mm (N = 4); holotype 
male 16 mm. 

Genitalia (Figs. 3-6). Uncus (Figs. 3, 4) long, slender, strongly curved apically. Valve 
flattened, evenly elongate and simple (Fig. 3). Anellus of aedeagus (Fig. 5) well defined 
and heavily armed with spicules. The everted vesica (Fig. 6) consists of two divergent 
lobes, one the ductus ejaculatorius, and the other a broad, shallow, evenly rounded blind 
pouch. On the side of the vesica below the ductus ejaculatorius is a small but well developed 
multitoothed cornutus. The blind pouch is densely clothed terminally with small cornuti. 
Below this cluster at the end of aedeagus is a second cluster of stouter cornuti. 

Female. Similar exteriorly to male but with the following differences. Palps porrect, 
about three times length of head. Body vestiture shorter, much less dense than that of 
male. Antemedial line more faint; less contrast in color of basal and medial areas. Sub- 
terminal dots more developed. Two paratype females paler and less contrasting, with 
grayish lavender scales overlying tan shade beyond postmedian line. Orbicular visible as 


VOLUME 41, NUMBER 2 105 





Fics. 1-7. Bomolocha appalachiensis. 1, Holotype male. 2, Allotype female. 3-7. 
Genitalia. 3, Holotype male, ventral view (aedeagus removed); 4, Holotype male, lateral 
view (aedeagus removed); 5, Holotype aedeagus, vesica not everted; 6, Balsam Co., North 
Carolina, paratype, aedeagus, vesica everted; 7, Allotype female. 


a black dot in some females. Forewing length: 15-16 mm (N = 7); allotype female 16 
mm. 

Genitalia as illustrated (Fig. 7). 

Diagnosis. Male genitalia of B. appalachiensis were compared with those of B. manalis 
(Walker), B. baltimoralis (Guenée), B. bijugalis (Walker), B. palparia (Walker), B. aba- 


106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lienalis (Walker), B. deceptalis (Walker), B. madefactalis (Guenée), B. sordidula (Grote) 
and B. edictalis (Walker), the other Bomolocha species from the Appalachian region. 
Features of uncus, basocostal margin of valve, and sclerotized structures of aedeagus are 
diagnostic. Uncus of palparia, madefactalis and sordidula most similar to that of appa- 
lachiensis in being relatively long, slender, and curved apically. 

_ Valve of appalachiensis simple. The basocostal corner on the inner face of the valve 
is more strongly protuberant or actually toothed in most other Bomolocha species ex- 
amined; most species also show a slightly more distal projection. In bijugalis the valve is 
nearly as simple as that of appalachiensis, but the basocostal area is stouter and more 
heavily ridged than in appalachiensis. 

Aedeagus of compared species of Bomolocha differs from that of appalachiensis in 
having clustered cornuti either more or less developed, and in lacking the multitoothed 
cornutus or having it well developed with an expanded base. 

In size and shape of wings, appalachiensis is similar to palparia and madefactalis. 
Shape of the postmedian line, and degree of contrast between the chocolate brown basal 
two-thirds and paler apical area easily distinguish appalachiensis. 

Types. Holotype. Male, Greenbrier State Forest, Greenbrier Co., West Virginia, 9 July 
1977 at UV, Linda Butler (Fig. 1). Allotype. Female, Triune, Monongalia Co., West 
Virginia, 22 July 1979 at UV, Linda Butler (Fig. 2). Paratypes. 2 females, same data as 
allotype except 16 and 17 June 1979; 1 female, Big Ugly Public Hunting Area, Lincoln 
Co., West Virginia, 28 June 1979, Linda Butler; 1 male, Balsam, Jackson Co., North 
Carolina, 17 July 1974, Douglas C. Ferguson, U.S. National Museum Slide No. 56126; 1 
male, 1 female, Oconee State Park, Oconee Co., South Carolina, 24 May 1978, Eric 
Quinter; 1 female, data as above except 15 August 1973; 1 female, Clemson, South 
Carolina, 26 May 1976, R. S. Peigler and J. W. McCord; and 1 male, Kingdom Come 
State Park, Harlan, Kentucky, 13 July 1984, Loran D. Gibson. 

Holotype and allotype in U.S. National Museum; paratypes in U.S. National Museum, 
West Virginia University Arthropod Collection, and collections of Eric L. Quinter, Bryant 
Mather, and Loran Gibson. 

Distribution. The localities given for the types represent the known distribution. All 
specimens were taken in the Southern Appalachian Mountains from Monongalia Co., 
West Virginia in the north to Oconee Co., South Carolina in the south. 

Comments. Fifteen species of Bomolocha were previously known from North America 
(Hodges 1983). Besides the nine species found in the Appalachian region (Forbes 1954) 
and compared with B. appalachiensis, other species are B. ramstadtii (Wyatt) described 
from Florida (Wyatt 1967), B. henloa Smith described from Arizona (Smith 1905), B. 
atomaria Smith described from South Dakota (Smith 1903), B. vega Smith described from 
New Mexico (Smith 1900), B. umbralis Smith described from Florida (Smith 1884), and 
B. variabilis (Druce) described from Central America (Druce 1890). Woodruff (1913) 
discussed his collection of four specimens of B. atomaria from Connecticut. 


ACKNOWLEDGMENTS 


I thank D. C. Ferguson (Systematic Entomology Laboratory, ARS, USDA, Washington, 
D.C.), E. L. Quinter (American Museum of Natural History, New York, New York), 
Bryant Mather (Clinton, Mississippi), and Loran Gibson (Taylor Mill, Kentucky) for 
loaning paratype specimens. I also thank Ferguson, J. W. Amrine (West Virginia Uni- 
versity) and J. E. Weaver (West Virginia University) for helpful comments on the manu- 
script. 

This paper is published with the approval of the West Virginia Agricultural and Forestry 
Experiment Station as Scientific Article No. 1945. 


LITERATURE CITED 


DrucE, H. 1890. Lepidoptera, Heterocera. In Godman, F. D. & O. Salvin. Biologia 
Centrali-Americana 1:345-440. 


VOLUME 41, NUMBER 2 107 


FORBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Part III 
Noctuidae. Cornell Univ. Agric. Exper. Sta. Mem. 329. 433 pp. 

HonpcEs, R. W. (ED.). 1983. Check list of the Lepidoptera of America North of Mexico. 
E. W. Classey Limited & The Wedge Entomological Research Foundation, London. 
284 pp. 

SMITH, J. B. 1884. An introduction to a classification of North American Lepidoptera. 
Bull. Brooklyn Entomol. Soc. 7:70-83. 

1900. A hundred new moths of the family Noctuidae. Proc. U.S. Nat. Mus. 

22(No. 1208):413-495. 

1903. New noctuids for 1908, No. 4, with notes on certain described species. 

Trans. Am. Entomol. Soc. 29:191-224. 

1905. New Noctuidae for 1905. Can. Entomol. 37:65-71. 

WoopruFF, L. B. 1913. Bomolocha atomaria Smith in Connecticut. Bull. Brooklyn 
Entomol. Soc. 8:46—47. 

Wyatt, A. K. 1967. A new Bomolocha from Florida (Noctuidae). J. Lepid. Soc. 21: 


125-126. 











Journal of the Lepidopterists’ Society 
41(2), 1987, 108-113 


THE TYPES AND STATUS OF PAPILIO TASSO STAUDINGER 


KURT JOHNSON 


Department of Entomology, American Museum of Natural History, 
Central Park West at 79th Street, New York, New York 10024 


AND 


DAVID MATUSIK 


Department of Entomology, Field Museum of Natural History, 
Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605 


ABSTRACT. Two syntypes, sole representatives of Papilio tasso, are critically ex- 
amined for the first time and a lectotype male designated. The previously unexamined 
male genitalia are described and illustrated as well as the hitherto unillustrated syntype 
female. Lectotype wing characters and distinctive symmetrical valval harpes strongly 
suggest tasso is a valid species of the torquatus group of Heraclides. However, it should 
be investigated whether the lectotype represents a hybrid of common H. torquatus Cramer 
and nearly extinct H. himeros Hoppfer, and the syntype female an aberration or hybrid 
of the H. polybius Swainson complex. 


Additional key words: taxonomy, Papilionidae, Heraclides tasso. 


Papilio tasso has been one of the most enigmatic of swallowtail 
butterflies. The only known specimens, the male and female syntypes 
at the Zoologisches Museum der Humboldt Universitat zu Berlin (ZMH), 
have remained unexamined by 20th century students of Papilionidae. 
Workers have either (a) retained tasso as a species based on Staudinger’s 
(1884 [1888]) description and figure of the male (Rothschild & Jordan 
1906, Jordan 1907, Munroe 1961, D’Almeida 1965, Hancock 1983), (b) 
questioned its status (K. S. Brown Jr. pers. comm.), or (c) suggested that 
the species might not exist (D’Abrera 1981). The tasso female has not 
previously been illustrated. 

All workers have associated tasso with the torquatus group of Her- 
aclides Hubner, including with it the following taxa (distributions from 
D’Abrera 1981, American Museum of Natural History [AMNH] and D. 
Matusik [DMC] collections): H. himeros (Hoppfer) (SE Brazil), H. tor- 
quatus (Cramer) (many subspecies from central Mexico to S-central 
South America), H. garleppi (Staudinger) (subspecies in W Amazon 
basin, Bolivia, and “Guianian region’), H. lamarchei (Staudinger) (Bo- 
livia and SE Brazil, N Argentina), H. hectorides (Esper) (SE Brazil W 
to Bolivia and S to N Argentina). Most of these species (except perhaps 
H. hectorides) are considered rare, and are poorly represented in col- 
lections. 

The sister group of the above taxa, the Heraclides anchisiades group, 
was recently reviewed (Johnson & Rozycki 1986). As part of an effort 
to clarify the status of terminal taxa in the torquatus group, we ex- 
amined the Papilio tasso syntypes. 


VOLUME 41, NUMBER 2 109 


eur tA SEO >, 


Coll. Sommer} 





Fic. 1. Papilio tasso. A, Lectotype male; B, Syntype female. Upper surfaces at left, 
under surfaces at right. 


Heraclides tasso (Staudinger) 
(Figs. 1-2) 


Papilio tasso Staudinger 1884 [1888]:19. 


Types. Lectotype male (Fig. 1A) (ZMH) labelled “Origin”, “Pap. tasso Stgr.’’, “Coll. 
Sommer’, “Zool. Mus. Berlin” (no locality label, two undecipherable labels); we attached 
the label “lectotype designated by K. Johnson and D. Matusik, 1987”; locality by original 
description— ‘Brazil’. Syntype female (Fig. 1B) (ZMH) labelled “Origin” “tasso female”, 
“Brasilia”, “Zool. Mus. Berlin” (two undecipherable labels). We designate the male as 
lectotype because precedent diagnostic studies utilize male genital characters, and in- 
complete collection data and other considerations leave doubt about conspecific association 
of the syntypes. 

Characters. The lectotype differs from all other yellow and black members of the 
torquatus group as follows: Wings. Both forewing surfaces completely lack yellow col- 
oration distad of median area; both hindwing surfaces lack basal black in yellow bands, 
broad yellow extending from median area completely to wing base; hindwing devoid of 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fic. 2. Male genitalia of the Heraclides torquatus group, valval harpe, inner lateral 
view, on left; socii, outer lateral view, above right; aedeagus, lateral view, on right. Locality 
of specimen listed first with number dissected in parentheses if different than 1, additional 
localities and numbers in brackets thereafter. A, hectorides, Santissima-Trinidad, Para- 
guay (2) [Montevideo, Uruguay]; B, lamarchei, topotype, Bueyes, Bolivia [Agua Blanca, 
Argentina; Ichilio, Bolivia]; C, garleppi garleppi, “Bolivia” [Yapacani, E. Bolivia]; D, g. 
interruptus (Staudinger), Sani Beni, Peru[Tingo Maria, Peru; Carabaya, Peru]; E, himeros, 
‘Mentor’, Brazil [Bahia, Brazil]; F, tasso, lectotype; G, torquatus torquatus, Buena Vista, 
Bolivia [Teffe, Brazil], terminus of harpe above aedeagus, t. polybius, Matto Grosso, Brazil; 
H, t. leptalea (Rothschild & Jordan), Balzapamba, Ecuador (2); I, t. orchamus (Boisduval), 
Muzo, Colombia [Cauca Valley, Colombia]; J, tolus tolus (Godman & Salvin) (species 
status sensu Beutalspacher & Howe 1984), Chiltapec (Oaxaca), Mexico [Cordoba (Vera 
Cruz), Mexico], terminus of harpe above aedeagus, tolus mazai Beutalspacher & Howe, 


VOLUME 41, NUMBER 2 i 


emphatic postmedian to submarginal markings, these instead limited on under surface 
to three elongate whitish teardrop-shaped postmedian markings with adjacent smaller 
spots, and on upper surface to three small yellow spots, cells CU1 to M2; hindwing tails 
bulbously ovate; forewing length: 43.0 mm. Genitalia. Valval harpes symmetrical, each 
an extremely thin shaft terminating in a large serrate-edged knob; central area of shaft 
constricted and bearing a large, ventral-pointing spine; aedeagus with markedly con- 
stricted terminus. Abdominal coloration. Yellow over entire lateral area dorsad to thin 
black dorsal stripe, with cephalad-pointing black incised marking over lateral area of 
seventh and eighth tergites. 


Status of H. tasso Based on Lectotype Male 


Characters of the male valval harpe distinguish various papilionid 
taxa (Munroe 1961, Hancock 1983). We also showed useful differences 
in some aedeagii (Johnson et al. 1985, Johnson & Rozycki 1986). Figure 
2 illustrates male genital characters of 14 taxa of the torquatus group. 
With the traditional species-level taxa are included eight taxa viewed 
by authors as subspecies of H. torquatus. Left and right H. tasso lec- 
totype valval harpes are symmetrical and, compared with other mem- 
bers of the torquatus group, are very distinctive. By traditional taxo- 
nomic criteria (Munroe 1961, D’Abrera 1981, Hancock 1983), wing 
and genitalic characters of the lectotype strongly suggest that H. tasso 
represents a valid species. If so, its long absence from collections may 
reflect extinction or, as with H. himeros, near extinction (Collins & 
Morris 1985). Some workers, however, suspect that certain distinctive 
and rare papilionids result from infrequent hybridization (K. S. Brown 
pers. comm.). If this is true, distributional data and combinations of 
wing and genitalic characters suggest that H. tasso might be a tor- 
quatus-himeros hybrid. Such hybridization might explain the fuzzy 
limbal wing pattern and long, terminally knobbed valval harpe of the 
lectotype. Other characters, however, including the lectotype’s sym- 
metrical valvae, central harpe spine, bulbous hindwing tails, and lack 
of apical yellow do not appear to be easily explained in this manner. 
In one example of a rare papilionid “species” representing an unusual 
phenotype of another (Johnson & Matusik 1987), the genitalia of the 
aberrant specimen were nondistinctive. 


Syntype Female of P. tasso 


Association of the male and female Papilio tasso syntypes is prob- 
lematic. As Staudinger noted, the markings of the female are compelling 
both as to uniqueness among torquatus group females and suggestion 


—_— 


Guerrero (Guerrero), Mexico; K, atsukaae Igarashi, San Salvador, El Salvador; L, tolmides 
Godman & Salvin, Costa Rica. Dissections in AMNH and DMC except tasso. The last 
two are taxa of uncertain status. 


BZ JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of affinity to the male syntype. Workers who suspect H. tasso to be a 
hybrid suggest the syntype female may be an aberration of the non- 
yellow H. torquatus polybius Swainson complex (K. S. Brown pers. 
comm.). The syntype female resembles the lectotype in the extent and 
location of the white median to basal bands and in the bold lateral 
streaks on the under surface of the discal cell. Examination of torquatus 
group females in AMNH, DMC, British Museum (Natural History) 
(BMNH), Carnegie Museum of Natural History, Field Museum of Nat- 
ural History, and Allyn Museum of Entomology indicates the syntype 
female is distinctive. Not only does it differ from congeners in the wing 
characters resembling the lectotype, but no other female specimen has 
extensive pink-orange on the hindwing or an abdomen almost com- 
pletely white (congeners have a variably wide white lateral stripe). This 
extent of abdominal white is compatible with the extensive lateral 
yellow distinctive of the tasso lectotype and, to a lesser extent, males 
of H. himeros. Among torquatus group females, widening of the white 
forewing bands sometimes occurs in H. torquatus and H. hectorides. 
Usually, however, this widening does not include the discal cell, which 
is profusely invaded by white in the tasso syntype female. We know 
of no other specimen with basal to median white on the hindwing. A 
specimen of uncertain identification in BMNH labelled “Rio de Janeiro, 
Brazil” is similar to the tasso syntype female in wing and abdominal 
color characters except that it lacks basal to median white on both 
hindwing surfaces. We have not been able to ascertain if this is the 
same BMNH female cited by Rothschild & Jordan (1906:622) as bearing 
Gray’s label “P. polybius ‘variation a’”’ and appearing to them as syn- 
onymous with H. tasso. Original labels from Gray’s curations apparently 
have not always survived. However, since the above female was seg- 
regated, it may be the same specimen. (We affixed a label to it, “H. 
tasso? ...”, referring to this paper.) 

At present, study of the H. tasso female has limited value since 
females of the torquatus group are so rare in collections that the samples 
noted above would allow only comparison with H. torquatus and H. 
hectorides. 


CONCLUSIONS 


Traditional taxonomic criteria strongly suggest the lectotype of H. 
tasso represents a valid species. If so, it is possibly extinct or perhaps 
has not been collected since its original description. A number of “rare” 
papilionid taxa are known from only a few specimens (D’Abrera 1981, 
Johnson et al. 1985, 1986a, 1986b, 1986c); others have been collected 
only in disparate time periods (Collins & Morris 1985, Johnson et al. 


VOLUME 41, NUMBER 2 113 


1985, 1986b). Field and biological work must determine if extant natural 
populations exhibit the phenotypes of the Papilio tasso syntypes and 
whether their unique characters are attributable to hybridization or 
aberration. 


ACKNOWLEDGMENTS 


We thank H. J. Hannemann (ZMH) for loan of the types, and K. S. Brown Jr. (Uni- 
versidade Estadual de Campinas, Sao Paulo, Brazil) for discussions. The following searched 
collections and answered queries: P. R. Ackery (London), Rienk de Jong (Leiden), D. L. 
Hancock (Bulawayo, Zimbabwe), O. H. H. Mielke (Curitiba, Brazil), L. D. Miller (Sarasota, 
Florida), Tommasso Racheli (Rome), J. E. Rawlins (Pittsburgh), R. K. Robbins (Wash- 
ington), Richard Vane-Wright (London), E. W. Schmidt-Mumm (Caracas, Venezuela). 
Two anonymous reviewers made helpful suggestions, F. H. Rindge (AMNH) and Eric 
Quinter kindly reviewed a draft, and L. F. Gall (Yale University) provided literature. 


LITERATURE CITED 


BEUTALSPACHER, C. R. & W. H. Howe. 1984. Mariposas de Mexico. Fasiculo I. Pa- 
pilionidae. La Preusa Medica Mexicana, S.A., Mexico City. xii + 128 pp. 

CoLLins, N. M. & M. G. Morris. 1985. Threatened swallowtail butterflies of the world. 
International Union for Conservation of Nature and Natural Resources, Cambridge, 
England. vii + 401 pp. 

D’ABRERA, B. 1981. Butterflies of the Neotropical realm. Part 1. Papilionidae and 
Pieridae. Landsdowne Editions, East Melbourne. 172 pp. 

D’ ALMEIDA, R. F. 1965. Catalogo dos Papilionidae americanos. Sociedade Brasileria de 
Entomologia, Sao Paulo. 366 pp. 

Hancock, D. 1983. Classification of the Papilionidae (Lepidoptera): A phylogenetic 
approach. Smithersia 2:1—48. 

JOHNSON, K. & D. Matusik. 1987. The status of “Papilio hipparchus”’ Staudinger 
(Papilionidae). J. Lepid. Soc. 41:65-69. 

JOHNSON, K., D. MATusIK & R. ROZYCKI. 1986a. A study of Protesilaus microdamas 
(Burmeister) and the little-known P. dospassosi (Riitimeyer) and P. huanucana (Varea 
de Luque) (Papilionidae). J. Res. Lepid. In press. 

JOHNSON, K. & R. Rozycki. 1986. A new species of the anchisiades group of Heraclides 
from Venezuela (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 94:383-393. 

JOHNSON, K., R. Rozycki & D. MaTusik. 1985. Species status and the hitherto unrec- 
ognized male of Papilio diaphora Staudinger (1891), (Lepidoptera: Papilionidae). J. 
New York Entomol. Soc. 93:99-109. 

1986b. Rediscovery and species status of the neotropical swallowtail butterfly 

Papilio illuminatus Niepelt (Lepidoptera: Papilionidae). J. New York Entomol. Soc. 

94:516-525. 

1986c. The female of Papilio xanthopleura Godman & Salvin. J. Lepid. Soc. 
40:65-66. 

JorDAN, K. 1907. Papilionidae, pp. 1-51. In Seitz, A. (ed.), Macrolepidoptera of the 
World. Vol. 5. Alfred Kernen Verlag, Stuttgart. vii + 592 pp. 

MuNROE, E. 1961. The classification of the Papilionidae (Lepidoptera). Can. Entomol. 
Suppl. 17. 51 pp. 

ROTHSCHILD, W. & K. JoRDAN. 1906. A revision of the American Papilios. Novit. Zool. 
13:412-752. 

STAUDINGER, O. 1884[1888]. Exotische Tagfalter, Theil I. In Staudinger, O. & E. Schatz 
(eds.), Exotische Schmetterlinge, Bd. 1, Bescreibungen, 333 pp, Bd. 2, Abbildungen, 
100 plts. 








Received for publication 6 October 1986; accepted 30 March 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 114-115 


GENERAL NOTES 


PUPAE OF EURYTIDES THYASTES AND OTHER 
LEPTOCIRCINE SWALLOWTAILS 


Additional key words: Papilionidae, Leptocircini, Neotropics. 


Immature stages of Neotropical kite swallowtails of the thyastes group (Rothschild & 
Jordan 1906, Novit. Zool. 13:726-734; Munroe 1961, Can. Entomol. Suppl. 17:17; Hancock 
1983, Smithersia 2:20) are undescribed. In 1982 I obtained from Herbert Miers of Joinville, 
Santa Catarina, Brazil, four living pupae of the local member of the group, Eurytides 
thyastes (Drury), which he had reared. He told me that pupae of this species are sexually 
dimorphic in color, with green females and brown males, and indeed there were two of 
each sex and the colors were so distributed. It thus seems probable that pupal color in 
this species is not affected by the pupation environment, but, unlike the many examples 
of environmentally-cued pupal color dimorphism in swallowtails (Hazel & West 1979, 
Ecol. Entomol. 4:393-400 and others), green and brown pupae of E. thyastes are both 
dark and probably cryptic on the same substrates. The natural pupation sites are not 
known. Eurytides thyastes has a disjunct distribution: the upper Amazon region from 
NE Ecuador to Bolivia, and in SE Brazil near the coast (Rothschild & Jordan 1906, above). 
In Brazil the larval food is reported by Miers to be Talauma ovata A. St.-Hil. (Magno- 
liaceae), locally called ‘baguact’. He reared the E. thyastes on leaves of this tree in 
October 1981. The species is univoltine in SE Brazil, and adults emerged in October 1982. 

Fig. 1 shows a female pupa. Length 29.5 mm, widest point 9.5 mm, width of last 
abdominal segment 3.3. mm; thoracic horn about 2 mm long, projecting at 30° to the 
axis of the body. Green (female) and brown (male) pupae differ as follows: female green 
on all areas except where there is black (thoracic horn, dorsal surface of thorax, ventrally 
and dorsally on abdominal segments 4-6), pale tan (over wing bases, midventral region 
of abdominal segments 7 and 8, and in front of thoracic horn) or white (subdorsal mark 
on each side of abdominal segment 1). Ventral surface of last abdominal segment is brown, 
as are small tubercles on abdominal segments. Brown male pupae have exactly the same 
distribution of black, pale tan and white, but the green of the female is completely 
replaced by dull gray-brown. Among described leptocircine pupae it resembles Eurytides 
epidaus (Doubleday) (Ross 1964, J. Res. Lepid. 3:9-17), especially the head and the blunt 
thoracic horn, but E. thyastes is broader in the last few abdominal segments. The edges 
of the last segment are thick and dark. The pupa would merge with a thick branch if 
attached to one, much as Papilio clytia L. apparently does. Igarashi (1979, Papilionidae 
and their early stages, Tokyo [in Japanese] I:98) says that P. clytia pupates on branches 
at least as thick as a finger (translation), and it, too, has a broad tip on the abdomen 
(Igarashi 1979, above, II:color plate 107). 

Both Munroe (1961, above) and Hancock (1983,-above) place E. epidaus in the marcellus 
group, but its pupa differs in most respects from that of E. marcellus (Cramer). The 
thoracic horn of E. marcellus is pointed, and the pupa overall is smooth and patterned 
like the underside of a leaf, resembling those of some Old World Graphium spp. (Igarashi 
1979, above). However, another member of the marcellus group, E. marcellinus (Dou- 
bleday), is, according to T. W. Turner (pers. comm.), “without any prominent projections’. 

In contrast, the lysithous group (asius of Hancock 1983, above) appears to be homo- 
geneous in having bulbous green pupae with a long thoracic horn that continues the dark 
line on the side of the thorax and abdomen. Four members of the group are known to 
have this character: E. belesis (Bates) (Ross 1964, above, fig. 4A, B); E. lysithous (Hubner), 
E. protodamas (Godart) (West, unpubl. obs., specimens in Zoology Department, Univer- 
sidade Federal do Parana, Curitiba, Parana, courtesy Prof. O. H. H. Mielke); E. asius 
(Fabricius) (H. Miers pers. comm.). 

Apparently, pupae of the protesilaus group are undescribed, but that of one member 
of the remaining neotropical leptocircine group (dolicaon group) was described, though 
not figured, by D’Almeida (1924, Ann. Soc. Entomol. France 93:23-30): “Pupa 33 mm 


VOLUME 41, NUMBER 2 LP5 





Fic. 1. Eurytides thyastes female pupa. Left to right: dorsal, left lateral, ventral 
views. 


long, 9 wide, conical, very elongate and slender towards the abdominal region, with a 
thoracic projection 7 mm long, triangular, horizontal, directed forwards and projecting 
over the cephalic region .. .” (D’Almeida 1924, above, D. A. West translation). Although 
sketchy, this description resembles something between Graphium doson (Felder) and G. 
agamemnon (L.) (Igarashi 1979, above, color plates 209, 213) and is quite different from 
E. thyastes. 

In two related respects the pupae of E. thyastes, E. marcellus and E. lysithous are 
alike: the sculpturing of the posterior margin of the pupal hindwing has no “tail”, and 
the developing adult tail within is folded back along the outer margin of the hindwing. 

Pupal form may be a useful additional character for sorting out relationships in the 
Papilionidae. The limited data suggest that the lysithous group is monophyletic but that 
the marcellus group may not be. The comparison of Old and New World Leptocircini 
shows Protographium, Lamproptera and Graphium (Igarashi 1979, above), among the 
former, to be most similar to E. marcellus and perhaps E. dolicaon, among the latter. 
Hancock (1983, above) places the thyastes and dolicaon groups closest to the Old World 
genera and the marcellus group more distantly. Pupal morphology does not unequivocally 
support those placements, but the pupae of few species of Neotropical Leptocircini have 
been described. 

I thank the colleagues named for observations and material help and Takehiro Asami 
for translating parts of Igarashi. Supported by a National Geographic Society Grant. 


Davip A. WEsT, Department of Biology, Virginia Polytechnic Institute and State 
University, Blacksburg, Virginia 24061. 


Received for publication 22 August 1986; accepted 3 February 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 116-117 


ROOSTING BEHAVIOR IN ADULT VANESSA CARDUI 
Additional key word: Nymphalidae. 


Nocturnal roosting behavior has been reported for many butterfly species, including 
Heliconius charitonia (Poulton, E. B. 1931, Proc. Roy. Entomol. Soc. London 6:71, and 
others), H. erato (Crane, J. 1955, Zoologica 40:167-197) and other species (Carpenter, G. 
D. H. 1931, Proc. Roy. Entomol. Soc. London 6:71; Clench, H. K. 1970, J. Lep. Soe. 24: 
117-120; McFarland, N. 1971, J. Lep. Soc. 25:144-145). Below I describe apparent court- 
ship activity and subsequent roosting for the night observed in Vanessa cardui L. 

After a year of almost total absence in the Kalispell, Montana area (only one specimen 
seen in 1985), V. cardui was abundant throughout the Flathead River valley and the 
surrounding mountains in 1986. 

In the Creston area E of Kalispell, the flood plain of the Flathead River consists of 
numerous sloughs and wetlands, many drained since the turn of the century. These are 
interspersed with remnant stands of spruce, and the sloughs are lined with cottonwood 
and other deciduous species. On 29 May 1986, Louis Nimeroff and I noticed large numbers 
of a medium-sized insect active around one cottonwood tree 18-19 m tall alongside the 
road, and about 30 m from any other tree. As it was shortly after 2100 h (MDT) and the 
sun had set, fading light did not allow us to identify the insects on the wing. Several 
attempts to net them failed, as they were flying rapidly about the tree, but eventually 
we succeeded, and found they were V. cardui. We stayed until flight ceased and available 
light was nearly gone. The insects seemed to be alighting on cottonwood leaves upon 
cessation of activity and roosting there for the night. We returned the following night 
for further observations. 

On that evening, 30 June, we arrived at 2044 h and noted that the V. cardui were 
already engaged in the same activity as the evening before. Individuals were “chasing” 
close in to the cottonwood tree. Groups of 2 to 8 individuals “danced”, flying rapidly in 
a circular motion around one another for 1 to 5 seconds, and then dispersed, each flying 
away in a different direction, although occasionally 2 or 3 would re-form into a new 
“dance” group and begin anew. This “dance” involved lateral as well as vertical dis- 
placement. At no time did any group fly vertically more than 4.5 m from the starting 
point before dispersing. This activity suggested courtship (Scott, J. A. 1985, The butterflies 
of North America, Stanford, California, p. 283) but attempts to net an entire group were 
not successful, and we were unable to determine sexual composition. At no time during 
the observation period, however, did we note mating. 

Samples were taken opportunistically; nine males and two females were collected. All 
were worn, and were apparently new migrants into the area, although no obvious mi- 
gration was noted. Fresh specimens did not appear in the area until mid-June. 

Flight activity was observed up and down the tree, from ground level to the top. It 
was virtually constant, with individual insects pausing to rest on any convenient surface 
(ground, fence post, leaf, tree trunk, observer) for no more than 5 seconds. At rest, the 
wings were often held open so that the rays of the setting sun would fall full on them, 
but after sunset, which occurred at 2115 h, wings were held closed, and some were 
vibrated briefly in a manner reminiscent of sphingid and saturniid preflight warm-up. 
This thermoregulatory behavior has been noted elsewhere (Ferris, C. D. & F. M. Brown, 
eds. 1981, The butterflies of the Rocky Mountain states, Norman, Oklahoma, p. 31; and 
Scott 1986, above, p. 41). After sunset, resting became more frequent and prolonged. 

Some individuals were noted chasing other flying insects that wandered into the area 
of activity, including a fly (Tabanidae) and a dragonfly. From time to time, an individual 
cardui would leave the area of the tree and fly into adjoining fields. Return activity was 
noted with about the same frequency, resulting in stability in the number of cardui active 
about the tree. 


VOLUME 41, NUMBER 2 117 


We checked the cottonwoods and other trees bordering the nearby slough, and noted 
V. cardui engaged in the same type of activity, but the insects were present at a lower 
density. The isolation of the cottonwood of our observations may well have acted to 
concentrate activity in its vicinity. 

Activity ended abruptly. At 2148 h, flight and “dance” activity was still high. Two 
minutes later, only a few individuals were noted flying, and by 2154 h, all flight had 
ceased. 

Individuals were observed flying into the tree and alighting on a leaf, where they 
remained at least as long as we could see them (until about 2200 h), and presumably 
passed the night there. All individuals observed (we estimate at least 100 insects present 
at any time) came to roost in the tree, but were widely dispersed, with no group roosting 
observed. We were unable to return to the site during the morning hours to observe 
dispersal for the day. 


RICHARD L. HARDESTY, 20 4th Avenue East, Apt. 20, Kalispell, Montana 59901. 


Received for publication 4 August 1986; accepted 6 February 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 117-118 


OVIPOSITION BY PARIDES ARCAS MYLOTES (BATES) 
(PAPILIONIDAE) ON A GRASS LEAF-BLADE 


Additional key words: Papilioninae, Troidini, Aristolochia, Acroceras zizanoides, 
Costa Rica. 


Neotropical Parides oviposit exclusively on Aristolochia (Aristolochiaceae) plants (Moss 
1919, Nov. Zool. 26:295-319; Cook, Frank & Brower 1971, Biotropica 3:17-20; Young 
1973, Psyche 80:1-21; 1977, J. Lepid. Soc. 31:100-108; Brown, Damman & Feeny 1980, 
J. Res. Lepid. 19:199-226). Here I report a single observation of Parides arcas mylotes 
Bates ovipositing on the leaf-blade of grass. 

At 1600 h on 28 July 1986, a “fresh” P. arcas mylotes was followed for ten minutes 
through the 10-20 cm high ground cover of grass and weeds in a small clearing within 
the cacao plantation “Finca Experimental La Lola” near Siquirres (10°06’N, 83°30’ W; 50 
m eley.), Limon Province, Costa Rica. The butterfly alighted on a 15 cm long Aristolochia 
sp. seedling partly concealed in dense grass, and placed one egg on a meristem leaf. The 
butterfly then flew around the vicinity of this vine, and returned a few minutes later and 
placed another egg on the leaf-blade of the grass Acroceras zizanoides (H.B.K.) Dandy 
(Graminae), about 25 cm from the Aristolochia (Fig. 1). 

I collected both the Aristolochia and the Acroceras zizanoides, placing them in a 
tightly-closed plastic bag, to rear the two eggs. Roots of both plants were placed in a 
water-pik in the bag. I wanted to determine if the larva from the leaf-blade egg would 
feed on the grass, move to the Aristolochia vine (closest leaves about 10 cm away), or 
become disoriented and not feed at all. For the following two weeks of observation, 
neither larva fed on Acroceras zizanoides. The larva from the egg on the grass crawled 
to the Aristolochia without attempting to feed on the grass. Both larvae fed on the 
Aristolochia. I tentatively conclude that the oviposition on the grass leaf-blade was an 
aberrant behavior. Because troidine butterflies such as Parides are specialist herbivores 
on Aristolochiaceae (Brower & Brower 1964, Zoologica 49:137—159; and others), it would 
be of interest to determine the frequency of such behavior in the wild. Given that the 
meristem leaf of Aristolochia and the leaf-blade of Acroceras zizanoides are both narrow 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





meristem leaves of Aristolochia sp., nonhost and host plants, respectively, for Parides 
arcas mylotes in Costa Rica. A second instar of P. arcas mylotes is on the ventral surface 
of the Aristolochia leaf. Bottom: Egg of P. arcas mylotes on the ventral surface of the 
grass leaf-blade. 


and similar in shape (Fig. 1), suggesting they are visually similar to ovipositing troidine 
butterflies as shown by Rausher and Papaj (1983, Anim. Behav. 31:341-347), a fresh, 
“inexperienced” butterfly, like the individual observed here, might visually confuse host 
and nonhost plants. Nonhost oviposition occurs in the troidine genus Battus when the 
nonhost is visually similar to the host (Papaj 1986, J. Lepid. Soc. 40:348-349). 

I thank D. R. Papaj for helpful comments and suggestions. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Received for publication 22 September 1986; accepted 12 February 1987. 


VOLUME 41, NUMBER 2 119 


Journal of the Lepidopterists’ Society 
41(2), 1987, 119-121 


MATING BEHAVIOR OF ACRAEA ANDROMACHA ANDROMACHA 
(FABRICIUS) (NYMPHALIDAE) IN NEW CALEDONIA 


Additional key words: Acraeinae, sphragis, courtship. 


Acraea andromacha andromacha (Fabricius) is reported from Australia (primarily N 
and E), the SW Pacific, and the Lesser Sunda Islands of Indonesia (Holloway, J. D. & J. 
V. Peters 1976, J. Nat. Hist. 10:273-318). It belongs to the mostly African subfamily 
Acraeinae (Nymphalidae) which feeds on Passifloraceae. The species is the only Austral- 
asian one of Acraea, and Passifloraceae occurs chiefly in the Neotropics and Africa 
(Heywood, V. H. 1978, Flowering plants of the world, Oxford Univ. Press, 335 pp.). 

During copulation, male Acraea secrete a sphragis or plug and deposit it externally, 
thus preventing multiple female matings (Eltringham, H. 1912, Trans. Roy. Entomol. 
Soc. London, pp. 1-374). A large sphragis is believed to prevent mating by blocking the 
release of pheromone (Eltringham, above) and by the male detecting the sphragis phys- 
ically (Scott, J. A. 1972, J. Res. Lepid. 11:99-127; for a review of the role of the sphragis, 
see Drummond, B. A. 1984, pp. 291-370 in Smith, R. L., Sperm competition and the 
evolution of animal mating systems, Academic Press). Common and Waterhouse (1972, 
Butterflies of Australia, Angus & Robertson, Sydney, 498 pp.) report the presence of the 
sphragis on Australian female A. A. andromacha. The sphragis is also known in other 
Acraeinae (Planema and Actinote), Papilionidae, and Danainae (Scott, above). 

Along with the sphragis, African species of Acraea and Planema and Parnassius spp. 
lack courtship rituals (Eltringham, Scott, above). Strong female pheromones are postulated 
to replace courtship for intraspecific recognition (Eltringham, Scott, above). This mating 
strategy (assuming the pheromones exist) may have evolved where congeneric species of 
similar appearance and courtship occur sympatrically, as in African Acraea. Because A. 
andromacha is geographically isolated from congeners, it is not faced with interspecific 
mating. 

This note confirms that A. a. andromacha has mating behavior typical of its congeners, 
presents evidence of multiple mating, and discusses possible gene flow effects resulting 
from single mating in an island environment. 

In New Caledonia, A. a. andromacha occurs commonly in open, dry habitats with 
secondary vegetation and Acacia scrub (Holloway & Peters, above). Its external appear- 
ance and fluttery slow-wingbeat flight are reminiscent of Parnassius species (Papilionidae). 
I observed mating of A. a. andromacha on 28 January 1984 5 km N of Nepoui on the 
W coast of New Caledonia. The site was dominated by Leucaena leucocephala (Lam.) 
Dewit, an introduced legume-shrub found throughout the Pacific that occurs naturally 
in tropical America. The Acraea and other butterflies (Junonia villida calybe [Godart], 
Anaphaeis java persithene [Boisduval]), and an arctiid moth (Utetheisa sp.) were nec- 
taring on Tridax procumbens L., a weedy roadside composite. At 1100 h, a male A. a. 
andromacha pounced on a female in flight, and seconds later, on the ground, climbed 
on her dorsum and copulated without a courtship dance (Figs. 1, 2). When disturbed, 
the pair flew, in copula, to a nearby Tridax flower. Although I did not observe which sex 
flew, the female presumably carried the male because she alighted on the flower. In some 
Acraea species, either sex flies, whereas in others the male or female flies (Scott, above). 

I found sphraga on 14 of 15 female specimens, 1 per specimen, collected by M. G. 
Pogue and me in New Caledonia; and on 10 of 12 in the U.S. National Museum from 
Australia (8), New Caledonia (2), and the New Hebrides (2 without sphraga). Spermato- 
phore counts were made on three New Caledonian specimens with sphraga and one 
without. Two of the three with sphraga showed evidence of multiple matings, with two 
partially dissolved spermatophores, while the remaining specimen with a sphragis had a 
single fully formed spermatophore. The specimen without a sphragis had no spermato- 
phores, indicating, along with the excellent condition of her wings, that she was virgin. 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1,2. Mating behavior and copulation in Acraea a. andromacha. 1, Male climbs 
on dorsum of female and without courtship begins copulation. Wings of male nearly 
closed, pointing toward viewer, right wings more visible than left. Female’s right hind 
wing positioned in front of her right forewing. 2, Male’s wings, now open, cover all but 
female's forewings, which are positioned beneath male’s hindwings. 


Multiple female matings, indicated by more than one spermatophore, have been reported 
in several other Acraea species (Owen, D. F. et al. 1973, Entomol. Scand. 4:155—-160). 
They may be explained by loss of the sphragis due in part to its solubility in water, or 
by a second mating while the sphragis is still soft (Drummond, above). 

Number of matings between individuals can affect gene flow between Lepidoptera 
populations. A female that mates only once, before migration, increases gene flow by 
transporting both her and her mate’s genes to a new environment (Scott, above). In 
contrast, a female that mates again after migration lessens gene flow because the latter 
male fathers her subsequent offspring (sperm precedence, Labine, P. A. 1966, Evolution 
20:580-586). 

Sphragal-induced monogamy is more common in mainland areas, and one would 
assume that it has less adaptive value on islands. Nonetheless, such monogamy may serve 
to replace genes removed by natural selection from isolated island populations. Mated 
female Acraea a. andromacha with persistent sphraga (preventing further matings) and 
vagility equal to unmated females would influence gene flow among populations on Pacific 
islands more than unmated females. However, occurrence of multiple spermatophores in 
Acraea reported above and in the literature does not support this hypothesis. 

My visit to New Caledonia was supported by the Smithsonian Institution; Department 
of Entomology, University of Minnesota; Dayton Natural History Fund (Bell Museum, 
University of Minnesota); and The Society of Sigma Xi. I thank M. G. Pogue for inviting 
me to accompany him on the expedition; D. R. Davis for arranging the expedition; Gordon 


VOLUME 41, NUMBER 2 LAL! 


McPherson of the Missouri Botanical Gardens for local arrangements in New Caledonia 
and identification of the plants; and J. A. Scott, and D. H. Clayton for critical comments 
on the manuscript. 


Marc E. EpstEIN, Department of Entomology, University of Minnesota, St. Paul, 
Minnesota 55108. 


Received for publication 12 March 1986; accepted 19 February 1987. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 121-122 


A MIGRATORY FLIGHT OF THE CALIFORNIA 
TORTOISE-SHELL BUTTERFLY 


Additional key words: Nymphalis californica, Nymphalidae. 


While migrations, swarms, and dense clusterings of butterflies are well documented, 
we believe any significant mass movement of a lepidopteran should be reported. Ulti- 
mately, published reports will form the basis for a clearer understanding of the conditions 
causing migrations. With such an objective, we here present field observations and data 
by one of us (RMK) of a unidirectional mass movement of Nymphalis californica (Bois- 
duval) (Nymphalidae), the California tortoise-shell butterfly. This insect is known as a 
“loner” or “singleton” (Ferris, C. D. & F. M. Brown 1980, Butterflies of the Rocky 
Mountain States, University of Oklahoma Press, Norman, 442 pp.), but also has been 
recorded in enormous numbers and as migrating (Ferris & Brown, above; Howe, W. H. 
1975, The butterflies of North America, Doubleday and Co., Garden City, New York, 
633 pp.; Williams, C. B. 1930, The migration of butterflies, Biol. Monogr. & Manuals No. 
IX., Edinburgh, Oliver & Boyd, London, 473 pp.). The observations were made in Cal- 
ifornia in July 1986. Specimens were identified by one of us (ENL). 

On 25 July 1986 while driving E on US Interstate Hwy. 80 (I-80) at Pla-Vada, which 
is on the boundary between Placer and Nevada counties, a dense flight of N. californica 
was noted moving to the SW. This locality is 12 km E of the junction of I-80 and California 
State Hwy. 20 at an elevation of 1860 m. For a road distance of 400 m, butterflies swarmed 
over the highway in such numbers that they hit the automobile faster than they could 
be counted. So many insects both living ard dead were in the air turbulence of automobiles 
that they constituted a distraction to motorists. The density of this moving population 
gradually decreased eastwardly for 1.5 km at which point no further butterflies were 
seen. The time was 1145 h (PDT), temperature was 19°C, and relative humidity 35% 
under clear skies. Wind was estimated to be at 10-12 kmh out of the N. 

About 15 minutes later, more eastwardly on I-80, at the Donner Summit Rest Area, 
Nevada Co., located 12 km W of the junction of California State Hwy. 89 (S) and 24 km 
E of Pla-Vada, at an elevation of 2203 m, another flight of the species was observed. This 
migration was as dense as that noted above, and presumably was a part of the same 
population surge. The most dense section of this portion of the swarm stretched about 
800 m along I-80. After the dense swarm was passed, lesser numbers were observed for 
8 km E of Donner Pass to the Donner Lake Interchange (elevation ca. 2000 m). This 
second encounter with what we assume to be the same flying population observed earlier 
in the day was flying SW under weather conditions similar to those noted above. 

At the Donner Summit Rest Area, 8 butterflies (2 6, 6 2) were collected from the dead 
in a windrow along the road. Dead butterflies numbered 20-50/m? of roadside area, and 
extended along I-80 for at least 3 km. No species other than N. californica were noted 
among the dead. While this estimate gives some idea of the large numbers killed, visual 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


estimates indicated that those in flight numbered 5-25 butterflies passing the migration 
corridor each second. The height of flight was 1-4 m above the ground. Repeated counts 
of this unidirectional flight gave similarly steady readings of the moving population. As 
judged from the abundance of dead insects compared with those passing in flight, this 
flight had been taking place for more than two h. 

Three days later, on 28 July 1986, on a westward return trip, the migration was still 
in progress, but with fewer individuals spread over a longer distance. On that date, 10- 
25 butterflies could be counted in the air from anywhere within a circle. This flight 
extended from the Donner Lake Interchange over the Donner Pass through Norden, Soda 
Springs, and Cisco to the junction of J-80 and California State Hwy. 20, a distance of 42 
km. Three specimens (2 6, 1 2) were collected at the W-bound Donner Summit Rest Area. 
Climatic conditions were clear skies, no wind, 18°C, and about 30% relative humidity. 
Migration was westward 1-9 m above the ground. All 11 voucher insects are in the Insect 
Collections of the Department of Entomology, Louisiana State University, Baton Rouge. 

Later, on 1 August 1986, at the Donner Summit Rest Area, no individuals were seen 
in flight. Apparently the migrating population noted four to seven days earlier had moved 
through, and was no longer to be seen here. 

We thank Joan Chapin, Curator, Louisiana State University Collections, Department 
of Entomology, and N. Knaus and Y. Thomas for technical support. 


RONALD M. KNAUS AND EDWARD N. LAMBREMONT, Nuclear Science Center, Louisiana 
State University, Baton Rouge, Louisiana 70803. 


Received for publication 16 September 1986; accepted 6 March 1987. 


VOLUME 41, NUMBER 2 123 


Journal of the Lepidopterists’ Society 
41(2), 1987, 123 


SPEYERIA COLLECTION OF PAUL GREY TO THE 
AMERICAN MUSEUM OF NATURAL HISTORY 


The incomparable collection of the genus Speyeria (Nymphalidae) of Paul Grey has 
been deposited in the American Museum of Natural History. It consists of some 19,700 
mounted specimens, including 263 paratypes, with series from throughout the range of 
every species. The number of specimens ranges from a low of 94 for diana up to 4366 
for atlantis. This material has been incorporated into the Museum’s collection, which 
includes the Gunder and dos Passos specimens, among others, to form a study collection 
of 37,800 butterflies in this one genus. 

In addition to the specimens, about 150 genitalic preparations were received. These 
were made to study and help define each species, and formed the basis for “A genitalic 
survey of Argynninae (Lepidoptera, Nymphalidae)” by C. F. dos Passos and L. P. Grey 
(1945, Am. Mus. Novitates, no. 1296, 29 pp., 54 figs.). Additional study led the same 
authors to their “Systematic catalogue of Speyeria (Lepidoptera, Nymphalidae) with 
designations of types and fixations of type localities” (1947, Am. Mus. Novitates, no. 1370, 
30 pp.). Grey made a list of the specimens and slides used in the former paper, which is 
in the museum’s files; not all the material studied was available in the Grey and dos Passos 
collections. 

A catalogue on 3” x 5” index cards has been presented to the Museum also, listing and 
commenting on many of the specimens, by species and locality, that were studied by 
Paul Grey over the years. This takes up about two linear meters. Much valuable distri- 
butional data is included in this file; it also lists many butterflies not in his collection. 

The Speyeria collection of the American Museum of Natural History now contains not 
only the primary types of about one-third of the approximately 160 available names, but 
paratype and topotypical material of all, or nearly all, the specific and infraspecific names. 
The study collection has the specimens arranged by counties within the contiguous United 
States, and by neighboring areas in Mexico, Canada, and Alaska. Study of the distribution 
and variation within each species is simplified when the butterflies are pinned out in this 
manner. One result of this curating is that it quickly became evident to me that the 
majority of the subspecific names proposed in this genus are, at best, but random points 
on or at the ends of various clines, and hence are of little or no scientific value. There 
appear to be very few completely allopatric populations to which legitimate names might 
be attached. 


FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural 
History, New York, New York 10024. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 124-125 


MANUSCRIPT REVIEWERS, 1986 


The merit of a scientific journal depends on the quality of its reviewers as well as its 
authors, but the former are usually unknown to readers. The Journal acknowledges with 
gratitude the services of the people listed below from whom the editor received manuscript 


reviews in 1986. 


Phillip R. Ackery, London, UK 

W. Lee Adair Jr., Temple Terrace, FL 
James K. Adams, Lawrence, KS 

Robert L. Allen, Mission Viejo, CA 
David A. Andow, St. Paul, MN 

Richard T. Arbogast, Savannah, GA 
Paul H. Arnaud Jr., San Francisco, CA 
Richard A. Arnold, Pleasant Hill, CA 
Andrew Atkins, Dudley, NSW, Australia 


R. Robin Baker, Manchester, UK 

Edward M. Barrows, Washington, DC 
May R. Berenbaum, Urbana, IL 

André Blanchard, Houston, TX 

Carol L. Boggs, Stanford, CA 

M. Deane Bowers, Cambridge, MA 

K. S. Brown Jr., Campinas, Sao Paulo, Brazil 
Peter F. Brussard, Bozeman, MT 

John M. Burns, Washington, DC 

Linda Butler, Morgantown, WV 


Thomas W. Carr, Whitehouse, OH 

Timothy D. Cary, Stony Brook, NY 

Frances S. Chew, Medford, MA 

J. F. Gates Clarke, Washington, DC 

Jan F. B. Common. Toowoomba, Queens- 
land, Australia 

Charles V. Covell Jr., Louisville, KY 

David F. Crosby, E. Melbourne, Victoria, 
Australia 


Robert P. Dana, St. Paul, MN 

Robert Dirig, Ithaca, NY 

Julian P. Donahue, Los Angeles, CA 
John C. Downey, Cedar Falls, IA 
Arnold T. Drooz, Olustee, FL 

Boyce A. Drummond III, Florissant, CO 


Paul R. Ehrlich, Stanford, CA 

John N. Eliot, Taunton, Somerset, UK 
Thomas F. Emmel, Gainesville, FL 
Marc E. Epstein, St. Paul, MN 
Edward W. Evans, Manhattan, KS 


Douglas C. Ferguson, Washington, DC 

Clifford D. Ferris, Laramie, WY 

H. A. Freeman, Garland, TX 

Timothy P. Friedlander, College Sta- 
tion, TX 


Edward V. Gage, San Antonio, TX 
Lawrence F. Gall, New Haven, CT 
George L. Godfrey, Champaign, IL 


Richard D. Goeden, Riverside, CA 

Gary G. Grant, Sault Ste. Marie, Ontario, 
Canada 

Nancy Greig, Austin, TX 


Dale H. Habeck, Gainesville, FL 

David L. Hancock, Bulawayo, Zimbabwe 

Peter Harris, Regina, Saskatchewan, Can- 
ada 

Paul Hendricks, Pullman, WA 

John B. Heppner, Gainesville, FL 

Ronald W. Hodges, Washington, DC 


Kurt Johnson, New York, NY 


Roy O. Kendall, San Antonio, TX 

Joel Kingsolver, Providence, RI 

Edward C. Knudson, Bellaire, TX 

Norbert G. Kondla, Lethbridge, Alberta, 
Canada 


J. Donald Lafontaine, Ottawa, Untario, 
Canada 

Robert C. Lederhouse, E. Lansing, MI 

Norman C. Leppla, Gainesville, FL 


R. H. T. Mattoni, Beverly Hills, CA 

Sterling O. Mattoon, Oroville, CA 

Javier de la Maza Elvira, Tlalpan, DF, Mex- 
ico 

Tim L. McCabe, Albany, NY 

W. C. McGuffin, Ottawa, Ontario, Canada 

James S. Miller, Washington, DC 

Lee D. Miller, Sarasota, FL 

Scott E. Miller, Honolulu, HI 

Thomas A. Miller, Frederick, MD 

Eugene Munroe, Dunrobin, Ontario, Can- 
ada 

Dennis D. Murphy, Stanford, CA 

Judith H. Myers, Vancouver, BC, Canada 


Christopher D. Nagano, Los Angeles, CA 
Ray B. Nagle, Tucson, AZ 
H. H. Neunzig, Raleigh, NC 


Paul A. Opler, Ft. Collins, CO 


Richard D. Peterson, Fargo, ND 
Michael J. Plagens, Tucson, AZ 
Jerry A. Powell, Berkeley, CA 
Robert M. Pyle, Gray’s River, WA 


Stuart J. Ramos, Mayaguez, PR 
John E. Rawlins, Pittsburgh, PA 
Frederick H. Rindge, New York, NY 


VOLUME 41, NUMBER 2 


Robert K. Robbins, Washington, DC 
Ronald L. Rutowski, Tempe, AZ 


Theodore D. Sargent, Amherst, MA 
Jack C. Schultz, University Park, PA 
Albert Schwartz, Miami, FL 

Dale F. Schweitzer, Boston, MA 
James A. Scott, Lakewood, CO 
Arthur M. Shapiro, Davis, CA 

Jon H. Shepard, Nelson, BC, Canada 
Ernest M. Shull, N. Manchester, IN 
Michael C. Singer, Austin, TX 


125 


Frank Slansky Jr., Gainesville, FL 
Frederick W. Stehr, E. Lansing, MI 


G. P. Waldbauer, Urbana, IL 

Thomas J. Walker, Gainesville, FL 
Raymond R. White, Palo Alto, CA 
Per-Olof Wickman, Stockholm, Sweden 
Ronald L. Wilkinson, Washington, DC 
Ernest H. Williams, Clinton, NY 


M. P. Zalucki, St. Lucia, Queensland, Aus- 
tralia 


Journal of the Lepidopterists’ Society 
41(2), 1987, 126-127 


BOOK REVIEWS 


CALIFORNIA BUTTERFLIES, by J. S. Garth and J. W. Tilden. 1986. University of California 
Press, Berkeley, California. 246 pp., 20 color plates, hardcover. $19.95. 


California leads the nation in lepidopterous interest according to membership in the 
Lepidopterists’ Society. This first of two contemporary books aimed at the California 
audience (Emmel & Emmel will be at least another year in process) is a great disap- 
pointment. The book offers little more and a great deal less than its classic 60-year-old 
antecedent, Comstock’s Butterflies of California. The rather out-of-perspective dust cover 
illustration is a metaphoric warning about the contents. Following on the heels of Opler 
and Krizek, and the just published Scott, this work falls in the category of an anachronism. 

The authors introduce the book as a field guide, so that “users may learn where to find 
the butterflies and skippers of California ... may learn how to observe these Lepidoptera, 
how to study their fascinating habits, how to record observations so they will have meaning 
to others, and how to collect and preserve specimens.’ Of these topics, the overwhelming 
emphasis is on collecting. With space at a premium, we here get a half page figure on 
how to mount a moth ball on a pin without mention of the Environmental Protection 
Agency listing of naphthalene as a potent carcinogen. In contrast, there are two and a half 
pages on disappearing butterflies. But not to despair. Even though major expansion of 
urban areas has led to major irreparable habitat destruction and extinctions, “freeways 
provide quick access to some more distant butterfly haunts.” I was unable to find anything 
substantive on how to study habits or make meaningful observations, although these two 
topics should charge the goals of today’s butterfly “collector.” 

The field guide aspect of the book is another unfilled promise. Here we are instructed 
to use the key to families and then thumb through the illustrations. Without dwelling on 
the quality of the latter, one must conclude this could be a very frustrating experience. 
There are no maps, only general distribution data, and limited information on flight 
periods. Life history data are exemplified for Glaucopsyche lygdamus: “egg echinoid, 
flattened, with a raised white network; larva bright green with a magenta dorsal band; 
pupa brown with black dots.” This kind of “description” is unfortunately not unique to 
this book, but stands as a sad commentary on the primitive and almost useless state of 
information on early-stage morphology. 

In the wisdom of recognizing that to look up information on, discuss, or exchange a 
specific butterfly, you need to know its name, Garth and Tilden than go on to unques- 
tioningly apply the 1983 Miller and Brown check list (as in Check list of the Lepidoptera 
of America north of Mexico, R. W. Hodges, ed.). Not to add more to this over-discussed 
issue, it may be worth noting that the decision will hardly help those attempting to“look 
up’ information on some of the appellations. Occidryas editha won’t yield much from 
Biological Abstracts, although this butterfly is widely used in population studies. A further 
deference to instability with the use of common names is incredible: “What should become 
of acommon name that is based on a scientific name that changes? The mountain vagabond 
was a perfect common name for the fritillary Argynnis (now Speyeria) montivaga. But 
because montivaga has been shown to be a synonym of egleis, its common name has 
been changed in the present field guide to Egleis Fritillary.”” Follow that logic and you 
might understand the charge of the Light Brigade. Failure of the authors to confront the 
ferment in taxonomy and nomenclature can be expected to turn off any serious users of 
the book, who, in consulting other resources will wonder what is going on, and why. 

A brief history of butterfly collecting in California was the section I personally found 
most interesting—but then I have lived a good part of it. Some of the really flavorful 
stories, those uniquely Californian, are missing. We should have recorded such events as 
the night an intense midget jumped up during a Lorquin club meeting and called John 
Comstock a Nazi murderer for taking 500 Atossa fritillaries in a day. Then, the midget 
may have had a point because Atossa was never common after that day and became 
extinct three decades later. 


VOLUME 41, NUMBER 2 127 


I must recommend waiting and hoping Emmel and Emmel will answer our prayers. 
It is difficult being so negative about a work that caps two lives of involvement by two 


fine gentlemen who have otherwise contributed a great deal. To do otherwise would not 
be fair to the present generation. 


R. H. T. MaTTONI, 9620 Heather Road, Beverly Hills, California 90210. 


Journal of the Lepidopterists’ Society 
41(2), 1987, 128-129 


BUTTERFLIES OF EUROPE, Vol. 8, Aspects of the Conservation of Butterflies in Europe, 
Otakar Kudrna, Ed. (and author). 1986. AULA-Verlag, Wiesbaden 1, West Germany. 
323 pp. Order from: E. J. Brill, P.O. Box 9000, 2300 PA Leiden, Netherlands. $129.50 


(series subscription price $111.25). 


Although Vol. 8 is last in the Butterflies of Europe series, it is the second volume 
published. The title belies the contents, which are broader than butterfly conservation. 
This book consists of a Preface, 6 chapters, and a 14-page Bibliography. There is no index, 
but there is a detailed table of contents. Chap. 1 is a 4-page introduction to the aims, 
scope, materials, and methods related to this treatise. Chap. 2 is also short (18 pp.) and 
makes a case for using native butterflies as environmental health indicator species. Several 
species-specific examples are presented. There is a discussion of the order Insecta, followed 
by description of the six main European biomes cited in the book. 

Chap. 3 (81 pp.) treats the material of the book’s title. Starting with a biogeographic 
history of European butterflies during the Holocene, the author progresses to anthropo- 
genic factors harmful to butterflies. The latter subject is treated in detail, and discussion 
is liberally illustrated with colored photographs of habitats. Kudrna has identified 11 
anthropogenic (introduced by man) and 4 natural factors that comprise the pressures on 
any butterfly species. Among the former are air pollution, agricultural activities, urban- 
ization, and overcollecting; among the latter, climatic conditions, parasitoids, competition, 
and predators. Each factor is discussed with appropriate photographs and recognition of 
sensitive species. Causes and effects are examined with suggestions for protection measures. 
The author's treatment appears well balanced; he does not come across as a zealot in any 
sense. A few traditional conservationists may be offended by some of Kudrna’s comments, 
but I believe he has researched his subject carefully and thoroughly. He has addressed 
butterfly collecting in great detail from the aspects of scientific collecting, hobby collecting, 
and commercial collecting. The treatment is fair and covers pros and cons equally. The 
author points out the value of scientific collecting for demographic and taxonomic re- 
search; that destructive effects of overcollecting are not directly comparable to the re- 
percussions of habitat destruction; and that in Germany (as an example) each species 
threatened by overcollecting is more seriously threatened by man-made pressures. These 
pressures are usually the main cause of decline in butterfly numbers, not collecting per 
se. He points out that rare butterflies are sought because of their rarity, and that commercial 
collecting may pose a threat to some. 

Kudrna is critical of “Red Data Books” and other lists of rare, threatened, or endangered 
species. He presents cogent and documented arguments to support his position. He also 
points out that even in Europe, where butterflies have probably been studied more 
intensively than anywhere else, there are vast gaps in the knowledge of geographic 
distributions of many species. He demonstrates that considerable misinformation regard- 
ing geographic distributions has been promulgated from author to author. The conse- 
quence is that butterflies have been listed as endangered in some areas where, in fact, 
there are no valid records that they ever occurred. He calls for objectivity and use of 
accurate data, rather than emotion and conjecture. This stance will be unpopular with 
some readers, but it is valid and the point needs to be made. The last portion of Chap. 
3 evaluates the present state of knowledge of European butterflies. Kudrna concludes that 
much has yet to be learned concerning ecology, faunistics, distribution, and early stages, 
a situation certainly not unique to Europe. 

Chap. 4 (103 pp.) is entitled “Applied taxonomy of European butterflies”. Nearly half 
of this chapter is devoted to interpretation and explanation of the Code of Zoological 
Nomenclature published by the International Commission on Zoological Nomenclature. 
This treatment is informative and useful, but one questions its inclusion in a volume 
dealing with butterfly conservation. The Code discussion is followed by a 14-page glossary 
of taxonomic terms. Again the information provided is useful, but out of place. It would 
be more appropriate for the yet-to-be published Vol. 2, Introduction to Lepidopterology. 
The remainder of Chap. 4 consists of an annotated checklist of European butterflies, a 
provisional synonymic checklist, and a 2-page summary of “priority tasks. in taxonomic 


VOLUME 41, NUMBER 2 129 


research’. Thus all of Chap. 4 would seem to belong in Vol. 2, although the chapter is 
certainly valuable and well presented. 

Chap. 5 (69 pp.) returns to conservation and is devoted to applied biogeography of 
European butterflies. Following a short prefatory section, it is composed primarily of 
maps and tabular data. The author first defines biogeography in a broad sense, as opposed 
to zoogeography and faunistics. The area of concern to the author is conservation of the 
natural environment using butterflies as a representative animal! group. Butterfly colonies 
across Europe are surveyed, with tabular distributional data for all species. In addition 
to qualitative data, Kudrna develops a Chorological Index (CI) for each nominal species 
based on the sum of numerical values for size, composition, and affinity of range (as 
defined in the text). The smaller the CI value, the more successful the species. Large CI 
values indicate endemic species in limited areas. Evaluations of the “health” at the species 
level of all European butterflies are tabulated. Health factors include population declines, 
habitat vulnerability, and species vulnerability. These factors are assigned numerical 
values, and Vulnerability Indices (VI) from 1 to 6 are calculated (6 indicating most 
vulnerable). The latter portion of Chap. 5, well illustrated by color photographs, discusses 
butterfly ecology and habitats. 

The final chapter (22 pp.) outlines a comprehensive program to conserve European 
butterflies. Certain sensitive species are identified and discussed, and color photographs 
of their typical habitats included. 

This book is clearly written and the author amassed and digested a formidable amount 
of data. He has attempted to quantify data related to biogeography and species vulner- 
ability, thus removing much of the conjecture associated with typical conservation trea- 
tises. Data are presented, conclusions drawn, and positive recommendations are made. 
As indicated by numerous photographs, the author visited and studied many regions in 
Europe and has first-hand information. 

Like its predecessor, this book is in English, is well manufactured and attractively 
produced. I noted a few typographical errors and lapses into German (such as ist for is). 
The only deficiency is the absence of a glossary of ecological terms. There is liberal use 
of arcane terms such as “nemoral’, “eurychoric’, ““eurytopic’’, “xerothermophils ’, ““ubi- 
quists’, etc., which will not be familiar to the general reader. Other terms appear to be 
literal translations of German compound words. Perhaps a glossary can be included in a 
future volume. 

Although restricted to European species, this book belongs on the shelf of any serious 
lepidopterist. It contains a wealth of information and procedures. The treatment is well 
balanced and well researched. A serious attempt was made to quantify information. It is 
a scientific approach, and I suspect may well serve as a model for subsequent books on 
invertebrate conservation in regions other than Europe. 


CLIFFORD D. FERRIS, Bioengineering Program, University of Wyoming, P.O. Box 
3295 University Station, Laramie, Wyoming 82071. 


Date of Issue (Vol. 41, No. 2): 26 June 1987 


\ 
Ba 


ia ot us: “hs vise Spa 


a tT , 
Aiaa0>alees a at 
wy) feats i 


‘ae at ¢ % 


es y: . | 
bcd a. 


tas ak alg? A 
x j \ tp ‘ise 4 4 
f 7 qyitt > a 









EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
ROBERT C. LEDERHOUSE, THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Categories of contributions are Articles, General Notes, Technical Com- 
ments, Book Reviews, and Obituaries. Short manuscripts concerning new state records, 
current events, and notices should be sent to the News, June Preston, Editor, 832 Sunset 
Drive, Lawrence, KS 66044 U.S.A. Journal contributors should prepare manuscripts 
according to the following instructions. 

Abstract: An informative rather than descriptive abstract should precede the text of 
Articles. 

Key Words: Up to five key words or terms not in the title should accompany Articles, 
General Notes, and Technical Comments. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely 
double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. 
Titles should be explicit and descriptive of content, including the family name of the 
subject, but must be as short as possible. The first mention of a plant or animal in the 
text should include the full scientific name with author. Measurements should be given 
in metric units; times should be given in terms of the 24-hour clock (0930 h, not 9:30 
AM). Underline only where italics are intended. 

Literature Cited: References in the text of articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the heading 
LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In General Notes, references should be given in the text as Sheppard (1961, Adv. Genet. 
10:165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1:23-30). 

Illustrations: Only half of symmetrical objects such as adults with wings spread should 
be illustrated, unless whole illustration is crucial. Photographs and drawings should be 
mounted on stiff, white backing, arranged in the desired format, allowing (with particular 
regard to lettering) for reduction to their final width (usually 4% inches). Illustrations 
larger than 8% x 11] inches are not acceptable and should be reduced photographically 
to that size or smaller. The author’s name, figure numbers as cited in the text, and an 
indication of the article’s title should be printed on the back of each mounted plate. 
Figures, both line drawings and halftones (photographs), should be numbered consecu- 
tively in Arabic numerals. The term “plate” should not be employed. Figure legends 
must be typewritten, double-spaced, on a separate sheet (not attached to the illustrations), 
headed EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of 
illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical rules as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the proofs. 
A page charge of $60 per full Journal page is applicable to all authors. Unfunded authors 
may apply to the editor for a Society page charge grant. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A 





CONTENTS 


ECOLOGICAL SIGNIFICANCE OF A POSTMATING DECLINE IN EGG 
VIABILITY IN THE TIGER SWALLOWTAIL. Robert C. Leder- 
house ¢: J. Mark Scriber 

DEMOGRAPHY OF THE UNSILVERED MORPH OF SPEYERIA MORMONIA 
IN COLORADO. Carol L. Boggs 

XANTHORHOE CLARKEATA (GEOMETRIDAE), A NEW SPECIES AND 
POSSIBLE ENDEMIC OF THE QUEEN CHARLOTTE ISLANDS, 


BRITISH COLUMBIA. Douglas C. Ferguson cccccccccceeenmnenne: 
A NEW SPECIES OF NEARCTIC BOMOLOCHA (NOCTUIDAE) FROM 
THE APPALACHIAN AREA. Linda Butler ee 
THE TYPES AND STATUS OF PAPILIO TASSO STAUDINGER. Kurt 
Johnson & David Matusik 00 0 


GENERAL NOTES. 


Pupae of Eurytides thyastes and other leptocircine swallowtails. David A. 
WEST een 


Roosting behavior in adult Vanessa cardui. Richard L. Hardesty 


Oviposition by Parides arcas mylotes (Bates) (Papilionidae) on a grass leaf- 
blade. Allen Mo Young 


Mating behavior of Acraea andromacha andromacha (Fabricius) (Nymphal- 
idae) in New Caledonia. Marc E. Epstein 0. 


A migratory flight of the California tortoise-shell butterfly. Ronald M. Knaus 
d> Edward N: Lambremont 0 
Speyeria collection of Paul Grey to the American Museum of Natural History. 
Frederick H. Rindge 


Manuscript reviewers, 1986 0000 


Book REVIEWS 
California butterflies sul 
Butterflies of Europe; Vol. 80 


Volume 41 1987 Number 3 


| ISSN 0024-0966 


JOURNAL 


_ LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


ae ee OC 


TO Wey 








7 October 1987 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


JERRY A. POWELL, President JEAN-FRANCOIS LANDRY, Vice 
DouGLAS C. FERGUSON, Immediate Past President 

President ATUHIRO SIBATANI, Vice 
JACQUELINE Y. MILLER, Vice President President 
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer 


Members at large: 


MIRNA M. CASAGRANDE M. DEANE BOWERS JULIAN P. DONAHUE 
EDWARD C. KNUDSON RICHARD L. BROWN JOHN E. RAWLINS 
FREDERICK W. STEHR PAUL A. OPLER Jo BREWER 


The object of the Lepidopterists’ Society, which was formed in May 1947 and for- 
mally constituted in December 1950, is “to promote the science of lepidopterology in 
all its branches, ....to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists Society, to: James P. Tuttle, Treasurer, 
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian 
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California 
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec- 
retary, 50 Cleaveland Rd., #3, Pleasant Hill, California 94523-3765, U.S.A. 


To obtain: 

Back issues of the Journal and News (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society’) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, California 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blvd., Los 
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and 
additional mailing offices. POSTMASTER: Send address changes to the-Lepidopterists’ 
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007- 
4057. 


Cover illustration: Semilooping larva of the strange noctuid Phyprosopus callitrichoides 
on Smilax. Sketch by Mark Klingler, Carnegie Museum of Natural History. Suggested 
by John E. Rawlins. 


VOUBRNAL OF 
Tue LeEPIDOPTERISTS’ SOCIETY 


Volume 41 1987 Number 3 


Journal of the Lepidopterists’ Society 
41(8), 1987, 131-140 


PATTERNS OF OVIPOSITION IN HEMILEUCA LUCINA 
(SATURNIIDAE) 


M. DEANE BOWERS 


Museum of Comparative Zoology, Harvard University, 
Cambridge, Massachusetts 02138 


AND 


NANCY E. STAMP 


Department of Biological Sciences, State University of New York, 
Binghamton, New York 13901 


ABSTRACT. Hemileuca lucina (Saturniidae) is a batch-layer, ovipositing on stems 
of Spiraea latifolia (Rosaceae) in the fall, three months after larval development is 
. completed. Eggs hatch the following spring. Patterns of oviposition and choice of ovi- 
position site were determined by sampling three natural populations of this moth in 
Massachusetts. Mean number of eggs per mass was 146, with no decrease in the weight 
of eggs laid later relative to earlier in the mass. Females made few mistakes in choice of 
hostplant, and chose stems in sunny locations near the edge of S. latifolia clumps. Egg 
masses were deposited near the ground at a mean height of 33 cm, and on twigs between 
1 and 5 mm diam. 


Additional key words: eggs, Spiraea latifolia. 


Choice of oviposition sites may be critical for the reproductive success 
of an individual insect. Although mistakes in the selection of appropriate 
oviposition substrates occur (Chew & Robbins 1984), generally females 
are discriminating in where they lay their eggs (Rausher 1979a, 1979b, 
Stamp 1982, Singer 1983, 1984, Williams 1983, Grossmueller & Le- 
derhouse 1985). Appropriate choices may be particularly important for 
insects that lay eggs in batches, because they may make only one to a 
few decisions of where to oviposit. This is in contrast to species that 
oviposit eggs singly, where females may make hundreds of such de- 
cisions. 

Most studies on oviposition behavior have dealt with species in which 
eggs are deposited the same season in which the larvae feed, when cues 
are available that indicate appropriate larval host plants and environ- 


132 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ment. For example, Rausher (1979a) demonstrated that ovipositing 
Battus philenor (L.) (Papilionidae), whose offspring feed in that same 
season, discriminate among host plants and benefit by choosing plants 
that will not senesce before the larval period ends. Williams (1981) 
found that ovipositing Euphydryas gillettii Barnes (Nymphalidae) but- 
terflies chose leaves of the host plant with an orientation towards the 
sun that minimized larval development time, a critical component of 
larval survival in the montane habitats with short growing seasons where 
these butterflies occur. In contrast, cues such as hostplant quality and 
microclimate, which are directly related to offspring survivorship, may 
not be available to species that deposit eggs in the growth season prior 
to that in which larvae feed. Some of these species deposit eggs a few 
weeks after the larval period [gypsy moth, Lymantria dispar (L.) (Ly- 
mantriidae), and tent caterpillars, Malacosoma americana (Fab.) and 
M. disstria (Hibn.) (Lasiocampidae)], whereas others oviposit months 
after the larval period (species of tribe Hemileucini in Saturniidae) 
(Ferguson 1971, Tuskes 1984). 

Our objective was to examine oviposition behavior and site selection 
of Hemileuca lucina Hy. Edw. (Saturniidae), an insect that lays batches 
of eggs three months after the larval period and in the growth season 
before that in which larvae feed. 

Like other hemileucines, H. lucina is univoltine. The flight period 
occurs in mid to late September and lasts about two weeks. Females 
eclose laden with eggs. They attract males by producing a pheromone, 
apparently as soon as their wings expand, as Tuskes (1984) noted for 
other hemileucines. In the laboratory (at about 22°C), copulation lasts 
1 to 2 h (x = 92 min + 50 SD, n = 13). Females begin depositing eggs 
shortly afterwards. 

Females deposit usually 1, but occasionally 2 (less than 5% deposit 
2), clusters of eggs in a ring around stems of the host plant, Spiraea 
latifolia (Ait.) Borkh. (Rosaceae) (Fig. 1). The female climbs onto a 
twig and curls the abdomen up and to the side around the twig for 
each egg deposited. She alternates from side to side, with two depositions 
made swinging the abdomen to the left and then two to the right. The 
first few eggs are laid in a sparse half-circle around the twig. The female 
then works her way up the twig, carefully inserting eggs into the gaps 
among the others, thus making a tightly packed egg ring. Usually, 
several rows are deposited, with the rows progressing up the stem. 

The eggs overwinter and, in Massachusetts, hatch sometime in May, 
depending on local weather conditions. Larvae are gregarious in the 
first four instars. Group size declines, and by the fourth or fifth instars 
solitary individuals are observed in the field (Cornell et al. 1987). Factors 
such as parasitoids and predators, weather, and food availability con- 


VOLUME 41, NUMBER 3 133 





Fic. 1. Egg mass of H. lucina. Top, longitudinal view. Bottom, stem of Spiraea latifolia 
cut to show arrangement of eggs around it. 


tribute to reduction in group size. Larvae are black with spines in the 
first five instars. The spines cause painful dermatitis when larvae are 
handled, particularly in the penultimate and ultimate instars. Larvae 
are solitary in the sixth instar and usually have a lateral yellow stripe 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and yellow mottling, but can be quite variable (D. F. Schweitzer pers. 
comm.). Although larval feeding in the early instars is confined to S. 
latifolia, we observed that later instars may feed on other plants, such 
as blackberry (Rubus sp.), cinquefoil (Potentilla sp.) and black cherry, 
Prunus serotina (Rosaceae). The larvae leave the host plant, often move 
some distance, and burrow into the soil to pupate, where they aestivate 
through the rest of the summer. 


MATERIALS AND METHODS 


Egg masses were collected from Dover, Norfolk Co., Massachusetts, 
in 1983 and 1984; Belmont, Norfolk Co., Massachusetts, in 1984; and 
Leverett, Franklin Co., Massachusetts, in 1984 and 1985. Number of 
eggs per mass was counted at these 3 sites. In 1985, we determined the 
percent of eggs hatching at Leverett. 

To examine whether the first eggs deposited weighed more than 
those deposited later, as has been found in other lepidopterans (Wel- 
lington 1965, Leonard 1974, Richards & Myers 1980, Jones et al. 1982, 
Wiklund & Persson 1983, Karlsson & Wiklund 1984, Harvey 1985), 
eggs from newly laid egg masses were detached from twigs and weighed 
individually. Because we were unable to determine the exact order of 
deposition but knew that eggs were laid from the bottom to the top, 
we removed nine eggs from the bottom ring and nine from the top 
ring of each of five egg masses. The null hypothesis was that no dif- 
ference in egg weight occurred between eggs laid first and those laid 
last. 

At Leverett in 1984 and 1985, we described the location of egg masses. 
We first measured the height at which they occurred on S. latifolia 
stems. On 19 May 1985, we measured, to the nearest 10 cm, their 
distance from the edge of S. latifolia clones. In addition, we noted 
whether masses were located in the sun or shade. We also searched 
outside dense clones of S. latifolia for egg masses. 

To determine what diameter of stem females selected for oviposition, 
and whether that reflected what was available, we compared diameter 
of stems containing egg masses with that of stems that were available. 
On 14 April 1985, the diameter of available stems was measured by 
establishing transects through clones of S. latifolia at Leverett. A tran- 
sect was set up through the middle of each of two clones, and another 
transect along the edge of these same clones, for a total of four transects. 
The height used for sampling diameter of available stems was based 
on mean height above ground of egg masses the previous year (1984) 
at that site (x = 24.9 cm + 11.4 SD, n = 83). Every stem within 5 cm 
of the transect was measured at a height of 25 cm. We assumed that 
relative availability of stems of different diameters did not change 


VOLUME 41, NUMBER 3 135 


TABLE 1. Number of eggs in egg masses of Hemileuca lucina from three sites in 
Massachusetts. Dover “a” refers to egg masses collected in 1983, hatching that year, and 
Dover “b” refers to egg mass remains collected in 1984, hatching in 1983. 


Location Year Mean + SD n 
Dover 1983a 156.4 + 44.1 7 
b 142.1 + 43.9 53 

1984 135.2 + 44.5 34 

Belmont 1982 SW Wey oyo MN 5 
1983 PANO esa tS) 2 

Leverett 1983 147.4 + 50.8 32 
1984 149.9 + 47.8 33 

Total 145.8 + 44.7 176 


during the study. New stems are produced each year, old ones grow, 
and there are thousands of stems available from which females may 
choose. 

Power of the statistical tests (probability of not committing a type II 
error, 1 — 6) was calculated as described by Cohen (1977). 


RESULTS AND DISCUSSION 


Description of the egg mass. A typical egg mass is shown in Fig. 1. 
Mean weight of an egg from a single mass was 1.63 mg + 0.16 SD and 
ranged from 1.13 to 1.89 mg (n = 118). Such variation in egg weight 
may or may not relate to larval fitness (Wellington 1965, Richards & 
Myers 1980, Wiklund & Karlsson 1984, Karlsson & Wiklund 1984, 
Harvey 1985). In general, weight of eggs when laid declines as lepi- 
dopteran females age (Wellington 1965, Leonard 1974, Richards & 
Myers 1980, Jones et al. 1982, Wiklund & Persson 1983, Karlsson & 
Wiklund 1984, Harvey 1977, 1985, Boggs 1986). Accordingly, for those 
species laying only one or two egg masses, egg weight would be pre- 
dicted to decline from first to last eggs laid in a single mass, as Wellington 
(1965) found for Malacosoma californicum pluviale (Dyar) (Lasiocam- 
pidae). However, we found that the first eggs deposited in an H. lucina 
egg mass were not significantly different in weight from eggs deposited 
last (Wilcoxon paired sample test, n = 5, P > 0.05). Because H. lucina 
females do not feed as adults, often mate and oviposit within a few 
hours of eclosion, and usually lay only a single egg mass, they may 
emerge with all eggs yolked and allocate resources to eggs randomly 
(Wiklund & Karlsson 1984). 

The mean number of eggs per mass was 145.8 (Table 1), and ranged 
from 42 to 235. Of 36 egg masses collected in Leverett in 1984, 2 did 
not hatch; of 33 egg masses collected there in 1985, again 2 failed to 
hatch. Even within an egg mass, not all eggs hatched. Of the 31 egg 


136 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NUMBER OF EGG MASSES 





Wh ji ya! Ns Lit Sorte —— = 
PANE SPEER Sisk Santen © Boeke NM ER Aa fis, Se RSs MIG Toy ender 2 Gg BER SERA GE 


O-10 11-20 2l-30 31-40 41-50 5I-60 6I-70 7I-80 81-90 91-100 


HEIGHT OF EGG MASS - CM 


Fic. 2. Distribution of above-ground height of Hemileuca lucina egg masses in 1984 
(n = 40) and 1985 (n = 53) at Leverett. 


masses from Leverett that hatched in 1985, a mean of 12.9% (+16.5 
SD, range 1.1-93.5%) of eggs per mass did not hatch. 

Location of egg masses. Females were quite specific about where 
they laid eggs. Of 38 egg masses at Leverett in 1985, only 2 were on 
nonhost plants, 1 on blackberry (Rubus sp., Rosaceae), and 1 on gold- 
enrod (Solidago sp., Asteraceae). Both ““mistakes’’ were located in the 
midst of dense clumps of Spiraea latifolia, and probably did not ad- 
versely affect larval survival. Newly hatched larvae may move more 
than 100 cm to find food. Mean height of egg masses at Leverett was 
32.8 cm (+17.3 SD, n = 98) (Fig. 2). Height distribution of masses did 
not differ significantly between 1984 and 1985 (x?, df = 6, P > 0.10, 
1 — B = 0.55, a = 0.05). Stem height of S. latifolia ranged up to about 
200 cm; thus, females were avoiding the higher sites. Eighty-five percent 
of the masses occurred below 50 cm (Fig. 2). 

Females chose stems of a particular diameter (Fig. 3). Egg masses 
occurred on stems between 1 and 5 mm diam., whereas the available 
stems ranged from less than 1 mm to 12 mm. The size categories of 


VOLUME 41, NUMBER 3 137 


NUMBER OF TWIGS 


USED - DOVER 
60 
n=98 
50 





USED.- LEVERE ET 
40 n=65 





AVAILABLE - LEVERETT 
40 n=I5l 





Eee 
AL sole S04 Se Gree Bs SION He tt 
TWIG DIAMETER - MM 


Fic. 8. Diameter of stems at height of 25 cm available to ovipositing Hemileuca 
lucina females compared to diameter of stems chosen for oviposition. 


stems used and those of available stems at Leverett were significantly 
different (x2, df = 4, P < 0.001) (Fig. 3). Diameter of stems chosen by 
females at Leverett and Dover were not significantly different (x°, df = 
4,P > 0.05, 1 — 6 = 0.57, a = 0.05) (Fig. 3). 

All egg masses occurred in portions of S. latifolia clones not shaded 
by trees or shrubs during the day (n = 38, Leverett 1985). Thus, females 
appeared to avoid shady locations. In addition, females chose oviposition 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PER CENT OF EGG MASSES 
5O 


40 










30 


20 


























a el 


£30 «= 3 | -6O 46/=SO ae 


DISTANCE OF EGG MASS 
FROM EDGE OF CLONE (cm) 


periphery —————————>_ interior 


Fic. 4. Distribution of egg mass position in clones of Spiraea latifolia at Leverett in 
1985. 


sites on the periphery (0-60 cm) of S. latifolia clumps (Fig. 4), where 
more sunlight was available. 

Our results show that H. lucina females chose stems of S. latifolia 
that were a specific subset of those available for oviposition. First, such 
stems were located near the periphery of a clump and in the sun. This 
may aid in synchronizing egg hatch with bud burst, which is important 
in other species (Feeny 1970, Williams 1981). Second, egg masses were 
low on the stem, which may keep at least some under snow during the 
winter and ensure a more even microclimate. However, mean maximal 
monthly snow depth was 21.7 cm (+20.7 SD; n = 102 months, 1971— 
80 for 2 sites, Blue Hill Observatory near Dover, and Worcester, Mas- 


VOLUME 41, NUMBER 3 139 


sachusetts; National Climatic Center 1971-80). That suggests that most 
egg masses (mean height 32.8 cm above ground) were seldom covered 
by snow. Third, females chose stems of a particular diameter. If stems 
are too small, the eggs may not fit together to make a tight ring, and 
if they are too large, too few rows may be deposited to secure the ring 
to the stem. Finally, mistakes in choice of hostplant species were rare 
and usually occurred in the midst of S. latifolia clumps, suggesting that 
females may be attracted visually or olfactorily to the host plant, and 
that contact with chemical cues from the stem is less important. 

Ovipositing females of other lepidopterans are reported to discrim- 
inate successfully among hostplant species (Chew & Robbins 1984), seek 
sunny locations (Williams 1981, Grossmueller & Lederhouse 1985), 
choose microhabitats where desiccation problems are lessened (Carroll 
et al. 1979), and confront physical constraints posed by host plants 
(Levin 1973). Female H. lucina exhibit all of these levels of discrimi- 
nation, even though they oviposit at a time when conditions may be 
different from those when larvae are feeding. 


ACKNOWLEDGMENTS 


We thank J. Cornell, C. Fonstad, P. Johnson, J. Glyphis, S$. Grunow, K. Hoy Burgess, 
Z. Larrin, G. Puttick, and T. Sargent for assistance in locating field populations of H. 
lucina, and with field and laboratory work. M.D.B. was supported by the Clark Fund of 
Harvard University and NSF grant BSR-8307353. 


LITERATURE CITED 


Boccs, C. L. 1986. Reproductive strategies of female butterflies: Variation in and 
constraints on fecundity. Ecol. Entomol. 11:7-15. 

Bowers, M. D. 1978. Over-wintering behavior in Euphydryas phaeton (Nymphalidae). 
J. Lepid. Soc. 32:282-288. 

CARROLL, M. R., M. T. WoosTER, W. H. KEARBY & D. C. ALLEN. 1979. Biological 
observations on three oak leaftiers: Psilocorsis quercicella, P. reflexella, and P. cryp- 
tolechiella in Massachusetts and Missouri. Ann. Entomol. Soc. Am. 72:441—447. 

CHEw, F. S. & R. K. ROBBINS. 1984. Egg-laying in butterflies. Symp. R. Entomol. Soc. 
Lond. 11:65-79. 

COHEN, J. 1977. Statistical power analysis for the behavioral sciences. Rev. ed. Academic 
Press, Orlando. 474 pp. 

CORNELL, J. C., N. E. Stamp & M. D. Bowers. 1987. Developmental change in 
aggregation, defense and escape behavior of buckmoth caterpillars, Hemileuca lucina 
(Saturniidae). Behav. Ecol. Sociobiol. 20:383-388. 

FEENY, P. P. 1970. Oak tannins and caterpillars. Ecology 51:565-581. 

FERGUSON, D. 1971. Bombycoidea: Saturniidae. Moths of North America. Fascicle 
20.2A. E. W. Classey Ltd., London. 153 pp. 

GROSSMUELLER, D. W. & R. C. LEDERHOUSE. 1985. Oviposition site selection: An aid 
to rapid growth and development in the tiger swallowtail butterfly, Papilio glaucus. 
Oecologia 66:68-73. 

HarVEY, G. T. 1977. Mean weight and rearing performance of successive egg clusters 
of eastern spruce budworm (Lepidoptera: Tortricidae). Can. Entomol. 109:487—496. 

1985. Egg weight as a factor in the overwintering survival of spruce budworm 

(Lepidoptera: Tortricidae) larvae. Can. Entomol. 117:1451-1461. 





140 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Jones, R. E., J. R. Hart & G. D. BULL. 1982. Temperature, size and egg production 
in the cabbage butterfly, Pieris rapae L. Aust. J. Zool. 30:223-232. 

KARLSSON, B. & C. WIKLUND. 1984. Egg weight variation and lack of correlation 
between egg weight and offspring fitness in the wall brown butterfly, Lasiommata 
megera. Oikos 43:376-385. 

LEONARD, D. 1974. Recent developments in ecology and control of the gypsy moth. 
Ann. Rev. Entomol. 19:197-229. 

LEVIN, D. A. 1973. The role of trichomes in plant defense. Quart. Rev. Biol. 48:3-15. 

RAUSHER, M. D. 1979a. Larval habitat suitability and oviposition preference in three 
related butterflies. Ecology 60:503-511. 

1979b. Egg recognition: Its advantage to a butterfly. Anim. Behay. 27:1034- 
1040. 

RICHARDS, L. J. & J. H. Myers. 1980. Maternal influences on size and emergence time 
of the cinnabar moth. Can. J. Zool. 58:1452-1457. 

SINGER, M. C. 1983. Determinants of multiple host use by a phytophagous insect 
population. Evolution 37:389-403. 

1984. Butterfly-hostplant relationships: Host quality, adult choice and larval 
success. Symp. R. Entomol. Soc. Lond. 11:81-88. 

STAMP, N. E. 1982. Selection of oviposition sites by the Baltimore checkerspot, Euphy- 
dryas phaeton (Nymphalidae). J. Lepid. Soc. 36:290-302. 

TuskEs, P. M. 1984. The biology and distribution of California Hemileucinae (Satur- 
niidae). J. Lepid. Soc. 38:281-309. 

WELLINGTON, W. G. 1965. Some maternal influences on progeny in the western tent 
caterpillar, Malacosoma pluwviale (Dyar). Can. Entomol. 97:1-14. 

WIKLUND, C. & B. KARLSSON. 1984. Egg size variation in satyrid butterflies: Adaptive 
vs. historical, “Bauplan’, and mechanistic explanations. Oikos 43:391—400. 

WIKLUND, C. & A. PERSSON. 1983. Fecundity, and the relation of egg weight variation 
to offspring fitness in the speckled wood butterfly Pararge aegeria, or why don't 
butterfly females lay more eggs? Oikos 40:53-63. 

WILLIAMS, E. H. 1981. Thermal influences on oviposition in the montane butterfly 
Euphydryas gillettii. Oecologia 50:342-346. 

WILLIAMS, K. S. 1983. The coevolution of Euphydryas chalcedona butterflies and their 
larval hostplants. III. Oviposition behavior and host plant quality. Oecologia 56:336- 
340. 








Received for publication 22 December 1986; accepted 8 May 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 141-144 


PREDATION BY ANOLIS LIZARDS ON BATTUS PHILENOR 
RAISES QUESTIONS ABOUT BUTTERFLY MIMICRY SYSTEMS 


FRANCOIS J. ODENDAAL, MARK D. RAUSHER, BETTY BENREY! 
AND JUAN NUNEZ-FARFAN! 


Zoology Department, Duke University, 
Durham, North Carolina 27703 


ABSTRACT. Anolis lizards in Texas make supposedly distasteful and poisonous Battus 
philenor adults a component of their natural diet. The lizards appear to suffer no ill 
effects, and individual lizards will eat Battus more than once. We followed individual 
female butterflies searching for oviposition sites for 90 h and observed 4 instances of 
predation in the field. We supplemented these observations with field experiments and 
a laboratory study. Results raise questions about the general importance of lizard predation 
in the evolution of butterfly mimicry systems. 


Additional key words: Papilionidae, aristolochic acid, distasteful butterflies, pipevine 
swallowtail. 


Mimicry is widespread in nature, and studies of predation on but- 
terflies have been prominent in the development of ideas about apo- 
sematic coloration (Brower 1958, Brower et al. 1963, Rothschild et al. 
1972). Most of these studies have involved examining the behavior of 
captive predators when offered palatable and distasteful butterflies. 
There is remarkably little information on predation of adult butterflies 
in nature, and published field observations deal almost exclusively with 
attacks of birds on butterflies (Fryer 1913, Rutowski 1978, Wourms & 
Wasserman 1985). Observations of natural predation by lizards are rare 
(Ehrlich & Ehrlich, 1982), yet “birds and lizards have long been con- 
sidered to be the major selective agents responsible for the extreme 
diversity of unpalatable and mimetic forms of butterflies in nature” 
(Boyden 1976). When wild Ameiva lizards in their natural habitat were 
fed live butterflies, they quickly became conditioned to avoid unpal- 
atable species (Boyden 1976). Ehrlich and Ehrlich (1982) observed 
iguanid lizards preying on tropical butterflies, and because different 
butterfly species seemed to be attacked differently, concluded that their 
observations supported the assumption that lizards are often strong 
selective agents in the evolution of butterfly color patterns and behavior. 

One classic study of mimicry in butterflies focused on the pipevine 
swallowtail and its mimics. The larvae feed on plants in the genus 
Aristolochia (Aristolochiaceae). The adults are distasteful to birds (Brower 
1958), presumably because they sequester distasteful aristolochic acids 
and related alkaloids, as do other Aristolochia-feeding papilionids (Euwe 
et al. 1968, Rothschild et al. 1972). These substances are poisonous to 


‘Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-233, Mexico 20, DF 


142 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


generalist insects in small quantities (Rausher 1979) and in vertebrates 
can cause acute renal failure (Hedwall 1961, Jackson et al. 1964). Mi- 
metic species include Papilio troilus, dark-form P. glaucus females, P. 
polyxenes, and Limenitis archippus archippus. 

During a field study of Battus philenor in the John Henry Kirby 
State Forest, Tyler Co., Texas, we observed individual females searching 
for oviposition sites for a total of 90 h from 22 March to 9 May 1985. 
We observed four instances of predation on Battus by lizards, as well 
as two unsuccessful attempts. These observations and additional ex- 
periments cause us to question whether qualities that render butterflies 
distasteful to birds also render them distasteful to lizards. 


Instances of Predation 


Our first observation of lizard predation involved an aging Battus female that alighted 
on a host plant and started to deposit eggs. Within seconds of alighting, a large Anolis 
pounced on her from a tree trunk about 0.4 m away, apparently killing her with its first 
bite, which covered her head and thorax. The lizard consumed the entire butterfly during 
the next 20 minutes. Getting the wings into its mouth appeared to be the most difficult 
part; it had to scrape the butterfly many times against a tree trunk using sideways 
movements of the head to work the wings in. Three other similar instances were also 
observed. Afterwards, we collected these lizards, and they exhibited no ill effects during 
24 h in captivity. 

In one case, a basking butterfly escaped when an Anolis pounced on it. Another time, 
a medium-sized Anolis jumped from a thin branch onto a stationary copulating pair on 
a twig about 30 cm below it, collided with them, dislodged the male, and fell about 2 m 
into the undergrowth below. Before its jump, the lizard was observed to climb slowly 
from near the pair up to the launching point. Apparently Anolis can perceive motionless 
Battus philenor. 


Reaction of Anolis to Offered Butterflies 


Tethered butterflies were presented to Anolis in the field. One female and five male 
Battus were allowed to fly past large (>15 cm) perching lizards. In all but one case in 
which the lizard appeared to be startled by the observer, the butterflies were seized 
immediately, sometimes by the body, and sometimes by the wings. The bodies but not 
the wings were eaten because tethers prevented lizards from freely scraping the wings 
against tree trunks. 

Two medium-sized (10-15 cm) lizards were each offered a Battus male. One lizard 
made no attempt to capture it. The second seized the butterfly immediately, but the 
tether became entangled in a twig; the lizard could not bring the dead butterfly to the 
ground and eventually abandoned it. We broke the wings of another butterfly near their 
base, making it unable to fly, and presented it to a large lizard perched on a tree trunk. 
The lizard immediately seized and consumed it entirely. 

Three small (<10 cm) lizards showed no interest in butterflies offered to them. They 
are almost certainly not large enough to capture and hold a Battus even if they tried. 
None of the lizards that ate butterflies exhibited any adverse symptoms during 24 h in 
captivity. 


Effect of Experience on Subsequent Predation 


Clearly, large Anolis lizards often kill and eat Battus philenor butterflies. It is possible 
that the predation we observed involved lizards that had not previously consumed a 
Battus adult. The question therefore remained whether eating one would discourage a 


VOLUME 41, NUMBER 38 143 


lizard from doing so again. To answer this question, we captured two large Anolis, placed 
them in a cage and fed both a Battus male the first day, a female the second day, and 
a male on each of the following two days. The lizards caught all butterflies immediately. 
Usually, the entire butterfly was consumed. We conclude that either Anolis lizards do 
not oe from experience to avoid Battus or the butterflies are not poisonous or distasteful 
to them. 


DISCUSSION 


Our observations indicate that Anolis lizards readily attack and con- 
sume an insect that serves as a model in a large mimicry complex. 
Qualities that render Battus distasteful to birds (Brower 1958) appar- 
ently do not render them so to Anolis. If this conclusion is applicable 
to other types of lizards, then lizard predation may have served less 
often than generally assumed as a major selection pressure causing the 
evolution of unpalatability or of mimicry. 

Three caveats must be added to this suggestion. First, because B. 
philenor is abundant in E Texas, it may constitute a potentially abun- 
dant resource for Anolis. It is thus plausible that these lizards have 
evolved to tolerate or detoxify the noxious compounds sequestered by 
B. philenor, and thus may not be representative of all lizards. Second, 
it is possible that in the year of our study alternative food resources for 
Anolis were scarce, and the lizards preyed on Battus despite distaste- 
fulness and possible subtle adverse effects. If so, then when alternative 
resources are more abundant, Anolis may exhibit less tendency to con- 
sume Battus. In such years the distasteful individuals would be pro- 
tected from lizard predation. This protection would favor the evolution 
of distastefulness and mimicry. Third, B. philenor in E Texas may not 
be distasteful, perhaps because it does not sequester noxious compounds. 
Two lines of evidence argue against this hypothesis: both butterflies 
and host plants contain aristolochic acids (Rausher unpubl. data); and 
female Papilio glaucus occur there predominantly in the black form, 
indicating that Batesian mimicry there is effective, which implies that 
Battus there is distasteful. 

Despite the caveats, we believe our observations suggest that lizards 
in our study have not been a major selective force in the evolution of 
mimicry and distastefulness in Battus philenor. Our results contrast 
sharply with those of Boyden (1976), indicating that the influence of 
lizard predation on the evolution of mimicry systems needs more in- 
vestigation. 


ACKNOWLEDGMENTS 
We gratefully acknowledge field assistance by R. C. Lederhouse, and permission from 


Gary Lacox, Texas Forest Service, to work in the Kirby State Forest. Two anonymous 
reviewers made useful comments. This research was funded by NSF grant BSR-8406870. 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


BOYDEN, T. C. 1976. Butterfly palatability and mimicry: Experiments with Ameiva 
lizards. Evolution 30:73-81. 

BROWER, J. V.Z. 1958. Experimental studies of mimicry in some North American 
butterflies. Part II. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus. 
Evolution 12:123-136. 

BROWER, L. P., J. V.Z. BROWER & C. T. COLLins. 1963. Experimental studies of 
mimicry. 7. Relative palatability and Mullerian mimicry among Neotropical butter- 
flies of the subfamily Heliconiinae. Zoologica 48:65-84. 

EHRLICH, P. R. & A. H. EHRLICH. 1982. Lizard predation on tropical butterflies. J. 
Lepid. Soc. 36:148-152. 

EuwE, J. VON, T. REICHSTEIN & M. ROTHSCHILD. 1968. Aristolochic acid I in the 
swallowtail butterfly Pachiloptera aristolochiae (Fabr.) (Papilionidae). Israel. J. Chem. 
6:659-670. 

FRYER, J. F. C. 1913. Field-observations on the enemies of butterflies in Ceylon. Proc. 
Zool. Soc. Lond. 2:613-619. 

HEDWALL, P. R. 1961. Einfluss der Aristolochiasaure auf die Nierenfunktionen von 
Ratten. Naunyn Schmiedeberg Arch. Exp. Path. 24:550. 

JACKSON, L., S. KOFMAN, A. WEISS & H. BRoDOvsKy. 1964. Aristolochic acid (NSC- 
50413): Phase I Clinical Study. Cancer Chemotherapy Rep. 42:35-37. 

RAUSHER, M. D. 1979. Coevolution in a simple plant-herbivore system. Ph.D. Thesis, 
Cornell University, Ithaca, New York. 240 pp. Diss. Abstr. Int. 39(11):527-B. Order 
No. 7910769. 

ROTHSCHILD, M., J. VON EUWE & T. REICHSTEIN. 1972. Aristolochic acids stored by 
Zerynthia polyxena. Insect Biochem. 2:334-348. 

RUTOWSKI, R. L. 1978. The courtship behaviour of the small sulphur butterfly, Eurema 
lisa (Lepidoptera: Pieridae). Anim. Behav. 26:892-903. 

Wouprms, M. K. & F. E. WASSERMAN. 1985. Bird predation on Lepidoptera and the 
reliability of beak-marks in determining predation pressure. J. Lepid. Soc. 39:239- 
261. 


Received for publication 1 August 1986; accepted 10 April 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 145-150 


NEW RECORDS OF BUTTERFLIES 
FROM THE WEST INDIES 


ALBERT SCHWARTZ 


Miami-Dade Community College, North Campus, 
Miami, Florida 33167 
Adjunct Curator, Florida State Museum, 
Gainesville, Florida 


FERNANDO L. GONZALEZ 
1825 W. 44th Place, Apt. 402, Hialeah, Florida 33012 


AND 


ROSE M. HENDERSON 


Milwaukee Public Museum, 800 West Wells St., 
Milwaukee, Wisconsin 53233 


ABSTRACT. 1[n338 person-days in the field, 503 specimens of butterflies were collected 
in the Cayman Islands and northern Lesser Antilles. Twenty-two species are reported for 
the first time from the islands of Anguilla, St.-Martin, St.-Barthélémy, Saba, St. Eustatius, 
and Nevis in the latter group (41 individual island records). Four species are reported 
for the first time from the Cayman Islands, as well as five new island records, bringing 
the number known from these islands to 44. 


Additional key words: distribution, Cayman Islands, Lesser Antilles. 


This paper deals with the Cayman Islands and the northern Lesser 
Antilles. It is based on two recent collections. Henderson collected on 
seven northern Lesser Antillean islands during 29 January—20 February 
1987 (23 person-days), visiting Anguilla, St.-Martin, St.-Barthélemy, 
Saba, St. Eustatius, St. Christopher (St. Kitts), and Nevis. Her butterflies, 
now in the collections of Schwartz (AS) and Gonzalez (FLG), consist 
of 230 specimens. A visit during 27 November-1 December 1985 (10 
person-days) to the Cayman Islands by Schwartz and Gonzalez resulted 
in 273 specimens. Voucher specimens of some species are in the Mil- 
waukee Public Museum. Nomenclature selectively follows Riley (1975), 
Miller and Brown (1981), and Brown and Heineman (1972). 


CAYMAN ISLANDS 


Gonzalez and Schwartz collected primarily on Grand Cayman, and 
Gonzalez spent one day on Cayman Brac; we did not visit Little Cay- 
man. Previous major contributions to knowledge of the Caymanian 
rhopaloceran fauna are Carpenter and Lewis (1943) and Askew (1980). 
Table 1 summarizes present knowledge of the fauna. Of the 44 Cay- 
manian species, 19 are now known from all three islands. Most species 
(41) are known from Grand Cayman (14 from only that island), with 
26 from Cayman Brac and 23 from Little Cayman. Inclusion of Her- 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


aclides machaonides (Esper) (Papilionidae) rests on D’Abrera (1981: 
32). But Carpenter and Lewis’s comprehensive work is based on the 
same collection, and they did not mention such an important record. 
Because the situation is anomalous, we do not include this species in 
the numbers on the Cayman Islands. Following are supplementary notes 
on the species new to these islands. 


Urbanus dorantes cramptoni Comstock (Hesperiidae). We collected 10 specimens (9 
AS, 1 FLG) on Grand Cayman, a first report. Our series was secured at three widely 
separated localities (Boatswain Bay, Cayman Kai, Old Man Bay); the broad distribution 
of this skipper on Grand Cayman suggests that it is not an extremely recent adventive, 
although it must have arrived since Askew (1980) collected there in 1975. 

Eurema elathea Cramer (Pieridae). This small-bodied species was recorded by Car- 
penter and Lewis (1943:377), and Askew (1980:128); both reported it only from Grand 
Cayman. Our four specimens (AS) from Cayman Brac are the first reported from that 
island; they were taken at the airport on the W end of the island. 

E. daira palmira (Poey) (Pieridae). First reported from Little Cayman and Cayman 
Brac by Askew (1980:129). Our 6 specimens (5 AS, 1 FLG) are from 4 localities on Grand 
Cayman (George Town, Cayman Kai, Old Man Bay, Boatswain Bay). 

Phoebis agarithe antillia Brown (Pieridae). “Authoritatively reported” but uncollected 
or unseen by Carpenter and Lewis (1943:372); record questioned by Askew (1980:1381). 
We have 4 specimens from Grand Cayman (Boatswain Bay; 3 AS, 1 FLG) and 1 from 
Cayman Brac (airport, W end; AS). 

Strymon martialis (Herrich-Schaffer) (Lycaenidae). Reported by both previous parties 
only from Little Cayman. We have 2 specimens (1 AS, 1 FLG) from Grand Cayman 
(Boatswain Bay) that constitute the first record for that island. 

S. columella cybirus (Hewitson) (Lycaenidae). Askew (1980:127) reported this hair- 
streak from Grand Cayman and Little Cayman. Gonzalez collected 2 specimens (1 AS, 
1 FLG) on Cayman Brac (airport, W end; Jennifer Bay), which constitute the first record 
for that island. 

Electrostrymon a. angelic (Hewitson) (Lycaenidae). First record for this species from 
the Cayman Islands. A single male (AS) collected. 

Hemiargus ceraunus filenus (Poey) (Lycaenidae). Known from Grand Cayman and 
Little Cayman (Carpenter & Lewis 1943:392; Askew 1980:128). Gonzalez took a single 
specimen (AS) on Cayman Brac (airport, W end). 

Junonia genoveva zonalis Felder & Felder (Nymphalidae). As on Jamaica (Turner & 
Parnell 1985), there are two species of Junonia on the Cayman Islands: J. genoveva and 
J. evarete. We took both on Grand Cayman (Boatswain Bay; 5 AS), but only J. g. zonalis 
on Cayman Brac (airport, W end; 1 AS, 1 FLG). On Grand Cayman, interspecific contacts 
were of common occurrence. The two species are readily distinguished on the wing, both 
by phenotype and behavior; J. evarete is more easily approached and captured. 

J. evarete (Cramer) (Nymphalidae). Widespread on Grand Cayman (George Town, 
Cayman Kai, Old Man Bay, Boatswain Bay; 8 AS, 3 FLG). 

Anaea cubana (Druce). First record of this cuban species from the Cayman Islands 
(Grand Cayman). We took 16 specimens (10 AS, 6 FLG) at 2 localities (George Town, 
Boatswain Bay). 


LESSER ANTILLES 


Butterflies of individual Lesser Antillean islands were studied in the 
late 19th and early 20th centuries. Godman and Salvin (1884, 1896) 
and Hall (1936) reported on the butterflies of Dominica, St. Vincent 
and Grenada, and St. Christopher. Later, Pearce (1969) and Schwartz 


VOLUME 41, NUMBER 3 147 


TABLE 1. Known distribution of the 46 taxa (44 species) of Rhopalocera on the Cayman 
Islands. 


Distribution 


Taxon Grand Cayman Little Cayman Cayman Brac 


Phocides pigmalion batabano xX 
Urbanus proteus domingo 

U. dorantes cramptoni 
Cymaenes t. tripunctus 
Hylephila p. phylea 

Calpodes ethlius 

Panoquina p. panoquinoides 
P. sylvicola 

Battus polydamas cubensis 
Heraclides aristodemus temenes 
H. a. tailori 

H. andraemon andraemon 

H. machaonides 

Ascia monuste eubotea 
Appias drusilla poeyi 

Nathalis iole 

Eurema elathea 

E. daira palmira 

Pyrisitia messalina 

P. lisa euterpe 

Abaeis nicippe 

Phoebis s. sennae 

P. agarithe antillia 

Aphrissa neleis 

Strymon martialis 

S. acis casasi 

S. a. gossei 

S. columella cybirus 
Electrostrymon a. angelia 
Hemiargus ammon erembis 
H. ceraunus filenus 

Leptotes cassius theonus 
Brephidium isophthalma thompsoni 
Agraulis vanillae insularis 
Dryas iulia 

Heliconius charitonius 
Junonia genoveva zonalis 

J. evarete 

Anartia jatrophae jamaicensis 
Siproeta stelenes insularis 
Phyciodes phaon 

Euptoieta hegesia 

Anaea cubana 

A. verticordia danieliana 
Danaus plexippus megalippe 
D. gilippus berenice 

D. eresimus tethys 

Total 23 26 


a 


~ Me OK 


bd 


mK Mm RK OM OK 


mA Mm MK MK 
MxM MM OK 


x MM KH RMR KM OK CK 


MK OS PK OS KS OS KO OO OM SS OO SK OO 
~ 
ees 


148 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and Jimenez (1982) surveyed the butterflies of Barbados and Montserrat, 
respectively. 

The only comprehensive work on the Lesser Antillean butterflies is 
that of Pinchon and Enrico (1969), the major source of Riley’s (1975) 
Lesser Antillean records. All distributions below are from these two 
sources. Henderson collected on most of the northern Lesser Antilles. 
Collection notes on selected species follow. 


Epargyreus z. zestos (Geyer) (Hesperiidae). Known from inner (St. Christopher) and 
outer (St.-Martin) chain islands and throughout the remainder of the Lesser Antilles to 
Grenada. One specimen (AS) taken on Saba, N of St. Christopher. 

Polygonus leo savigny (Latreille) (Hesperiidae). Previously known only from the north- 
ern islands of St. Christopher and Montserrat. Collected on St.-Barthélemy (2 FLG, 2 
AS). This is not only a northern extension in known range and a new island record, but 
the first record for the outer chain. 

Urbanus proteus domingo (Scudder) (Hesperiidae). Widespread from Saba to Grenada. 
Not previously reported from St. Eustatius (1 AS), Nevis (1 FLG, 1 AS), or St.-Barthelemy 
(1 FLG, 1 AS). . 

U. obscurus (Hewitson) (Hesperiidae). Widespread from Saba and Antigua S to Grenada 
and Barbados. Not previously reported from Nevis, whence we have 2 specimens (1 FLG, 
1 AS). 

Pyrgus oileus (Linnaeus) (Hesperiidae). Three specimens (1 FLG, 2 AS) from Nevis 
fill in the previous gap between St. Christopher in the north and Montserrat in the south. 

Wallengrenia ophites (Mabille) (Hesperiidae). Reported throughout the Lesser Antilles, 
from St.-Martin and St.-Barthélémy to St. Vincent. New records are St. Eustatius (1 AS), 
which lies between Saba and St. Christopher, and Nevis (1 AS) S of St. Christopher, 
whence W. ophites was known. 

Hylephila phylea phylea (Drury) (Hesperiidae). Widely distributed from St.-Martin 
and St.-Barthélémy to Grenada and Barbados; not previously reported from St. Eustatius 
(1 AS) or Saba (1 AS) N of St. Christopher, the northernmost inner-chain island whence 
H. p. phylea has been reported. 

Ascia monuste virginia (Godart) (Pieridae). Ranging from St.-Martin and Saba in the 
north to St. Lucia in the south. Previously unreported from Anguilla (1 FLG, 2 AS), St. 
Eustatius (2 FLG, 2 AS), and Nevis (1 AS). 

Pyrisitia lisa euterpe (Ménétriés) (Pieridae). Widespread from St. Christopher and St.- 
Martin in the north to St. Lucia and Barbados in the south. Not reported from Anguilla 
(2 FLG, 2 AS), Saba (2 FLG, 2 AS), St. Eustatius (2 FLG, 2 AS) or Nevis (1 FLG, 2 AS). 
The first two islands are northern extensions in known range within both chains. 

Eurema elathea (Cramer) (Pieridae). Broadly distributed from St.-Martin and St. Chris- 
topher to St. Lucia and Barbados. Not previously reported from Anguilla (8 FLG, 3 AS) 
N of St.-Martin. 

Phoebis s. sennae (Linnaeus) (Pieridae). Although broadly distributed throughout the 
West Indies, including the Lesser Antilles, where it has been reported from St.-Martin 
and Saba in the north to Grenada and Barbados in the south, there are no published 
records for Anguilla (1 AS), to the N of St.-Martin. In the inner chain, we have specimens 
from St. Eustatius (1 AS) and Nevis (2 FLG, 1 AS), which fill the previous gap between 
Saba and Montserrat. 

P. trite watsoni Brown (Pieridae). The only island known to harbor this species in the 
north is St. Christopher; on the more southern islands, it has been reported on Montserrat, 
Guadeloupe, Dominica, and St. Lucia. A single individual (AS) from Saba extends the 
known range from St. Christopher to the northernmost of the inner-chain islands. 

Strymon bubastus ponce Comstock & Huntington (Lycaenidae). Reported from St.- 
Barthélemy and St. Christopher in the north to Grenada in the south. Its range extends 
as far N as Saba, whence we have 1 specimen (AS), and Anguilla, where Henderson took 
6 (3 FLG, 3 AS). 


VOLUME 41, NUMBER 3 149 


Electrostrymon angerona (Godman & Salvin) (Lycaenidae). Occurs only on the inner- 
chain islands, from St. Christopher to Grenada. We have 1 specimen (AS) from Saba, 
thus “completing” its distribution on these northern islands. 

Leptotes cassius chadwicki Comstock & Huntington (Lycaenidae). Widely distributed 
as far S as Grenada, records fill in the northern gaps in both the inner (Saba [5 FLG, 8 
AS], St. Eustatius [2 FLG, 4 AS], Nevis [1 FLG, 1 AS]) and outer (Anguilla [2 FLG, 3 AS]) 
chains. 

Hemiargus thomasi woodruffi Comstock & Huntington (Lycaenidae). Previously known 
only from four Lesser Antillean islands, as well as Désirade. A single specimen (AS) from 
Nevis is a new island record, S of all other northern Lesser Antillean records. 

Agraulis vanillae insularis Maynard (Heliconiidae). Although known from four outer- 
chain northern islands, reported only from St. Christopher in the inner chain. We have 
specimens from Saba (2 FLG, 2 AS) and St. Eustatius (2 FLG, 2 AS) confirming its 
occurrence on the northernmost of the inner-chain islands. 

Junonia genoveva michaelesi Munroe (Nymphalidae). Widespread but unreported 
from Saba (1 FLG, 2 AS), St. Eustatius (1 AS), and Nevis (1 FLG, 1 AS) from the inner 
chain, and Anguilla (1 FLG, 2 AS) from the outer chain. 

Anartia j. jatrophae (Johansson) (Nymphalidae). Widely distributed as far S as Grenada 
and Barbados, but unreported from Nevis (2 FLG, 2 AS). Remarkably, Henderson did 
not see this species on any other island, although it was moderately abundant on Nevis. 
Known only from Antigua in the outer-chain islands. 

Biblis h. hyperia (Cramer) (Nymphalidae). In the northern islands, known only from 
St. Eustatius (where Henderson found it abundant on The Quill) and St. Christopher; 
Henderson took one specimen (AS) on Saba. 

Antillea p. pelops (Drury) (Nymphalidae). Reported only from St. Christopher and 
Montserrat. We have one specimen from Nevis (AS), between the foregoing islands. 

Anaea minor Hall (Apaturidae). Described from St. Christopher, reported from Antigua 
and Montserrat, as well as Guadeloupe. A specimen from Nevis (AS) fills in the gap 
between Montserrat and St. Christopher. Although Pinchon and Enrico (1969) reported 
these leaf butterflies from St.-Barthélémy, they stated that the population there tended 
toward the larger Puerto Rican A. borinquenalis Johnson & Comstock. 


Although the above island records add no new species to the 74 (95 
taxa) generally accepted from the Lesser Antilles, they add information 
on the distribution of 22 species on the northern Lesser Antilles. 


LITERATURE CITED 


ASKEw, R. R. 1980. The butterfly (Lepidoptera, Rhopalocera) fauna of the Cayman 
Islands. Atoll Res. Bull. 241:121-138. 

BROWN, F. M. & B. HEINEMAN. 1972. Jamaica and its butterflies. E. W. Classey, London. 
478 pp. 

CARPENTER, G. D. H. & C. B. Lewis. 1943. A collection of Lepidoptera (Rhopalocera) 
from the Cayman Islands. Ann. Carnegie Mus. Nat. Hist. 9:371—396. 

D’ABRERA, B. 1981. Butterflies of the Neotropical region. Pt. I. Papilionidae and Pier- 
idae. Lansdowne Editions, East Melbourne. xvi + 172 pp. 
GopmaNn, F. D. & O. SALVIN. 1884. A list of the Rhopalocera collected by Mr. G. 
French Angas in the island of Dominica. Proc. Zool. Soc. London 1884:314-320. 
1896. On the butterflies of St. Vincent, Grenada and the adjoining islands of 
the West Indies. Proc. Zool. Soc. London 1896:513-520. 

HALL, A. 1936. The butterflies of St. Kitts. Entomol. 69:274—278. 

MILLER, L. D. & F. M. BRowN. 1981. A catalogue/checklist of the butterflies of America 
north of Mexico. Lepidopterists’ Society, Los Angeles. 280 pp. 

PEARCE, E. J. 1969. The butterflies of Barbados. J. Barbados Mus. 33:76. 

PINCHON, PERE R. & P. ENRICO. 1969. Faune des Antilles frangaises. Les Papillons. M. 
M. Ozanne et Cie., Fort-de-France, Martinique. 258 pp. 





150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


RILEY, N. D. 1975. A field guide to the butterflies of the West Indies. New York Times 
Book Co., New York. 224 pp. 

SCHWARTZ, A. & C. J. JIMENEZ. 1982. The butterflies of Montserrat, West Indies. Bull. 
Allyn Mus. Entom. 66:1-18. 

TURNER, T. W. & J. R. PARNELL. 1985. The identification of two species of Junonia 
Hubner (Lepidoptera: Nymphalidae): J. evarete and J. genoveva in Jamaica. J. Res. 
Lepid. 24:142-153. 


Received for publication 23 May 1986; accepted 9 June 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 151-153 


A NEW SPECIES OF GRETCHENA (TORTRICIDAE) 
INJURIOUS TO PLANTED NEOTROPICAL WALNUT 


WILLIAM E.. MILLER 


Department of Entomology, University of Minnesota, 
St. Paul, Minnesota 55108 


ABSTRACT. Gretchena garai is described from four males and three females reared 
from shoots of Juglans neotropica Diels. (Juglandaceae) near Loja, Ecuador. Adults differ 
from all known congeners by their greenish vestiture and by absence of setae on the anal 
extensions of male cuculli. This is the first report of Gretchena from continental South 
America and the Neotropics. 


Additional key words: Olethreutinae, Eucosmini, Juglans neotropica, Ecuador. 


I describe this new species now because of its economic importance. 
It was discovered injuring leader shoots of Neotropical walnut, Juglans 
neotropica Diels., in Ecuador. One- to four-year-old plantings have 
been attacked repeatedly in the Loja Province highlands. Injured trees 
develop multiple leaders which lessen their chances of producing com- 
mercial timber. Walnut is important in Ecuadorian forestry, yielding 
timber similar in properties and value to North American black walnut, 
J. nigra L. Several species of Juglans, locally known as nogal, occur 
naturally from S Mexico through Central America and the cordilleras 
of Columbia, Ecuador, and Peru to Argentina (Chudnoff 1984). 

Gretchena, a genus of Eucosmini, currently comprises 11 Nearctic 
species, including the pecan bud moth, G. bolliana (Slingerland), and 
G. concitatricana (Heinrich), a black walnut shoot moth (Blanchard & 
Knudson 1983, Brown 1982, Naughton 1970, Powell 1983). Except for 
G. watchungana (Kearfott), whose larvae feed on Alnus (Betulaceae), 
the previously known larvae of Gretchena, numbering four species, 
feed on Juglandaceae (Miller 1987, Naughton 1970). Gretchena has not 
previously been reported from continental South America or the Neo- 
tropics (Powell & Razowski in press). 

I deemed the Ecuadorian Gretchena to be new after comparing 
specimens with descriptions of all check-list species of generically un- 
placed Eucosmini (Powell & Razowski in press). In the description 
below, character states that place the species in Gretchena (Brown 1982) 
are italicized. 


Gretchena garai, new species 


(Figs. 1-3) 


Male. Forewing 7.0-7.5 mm long (4n). Head. Length of second labial palpus segment 
1% eye diameter, whitish except for brownish black near base, middle, and apex; third 
segment % length of second. Face white, vertex yellowish. Thorax. Whitish, tegula white, 
brown, and green. Forewing termen concave, costal fold extending from base to middle 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-3. Gretchena garai. 1, Wings of holotype female; 2, Male genitalia (prep. 
WEM 2110862); 3, Female genitalia (prep. WEM 211086). 


and enclosing a white hair pencil, upper side patterned as in Fig. 1, basal patch more 
distinct on inner margin than on costa, geminations on costa white, darkest appearing 
areas brownish black, the less dark green, underside brownish gray, fringe brownish black. 
Hindwing upper and undersides, including fringe, brownish gray. Abdomen. Brownish 
gray. Genitalia (Fig. 2) (4n): Uncus absent; socii separate, bent dorsally just beyond 
middle; aedeagus % as long as valva, supported by long juxtal caulis, beaked at apex, 
vesica with 20-30 tightly packed cornuti % as long as aedeagus; valval neck sparsely 
spined, cucullus drawn out at lower anal margin to a spinelike tip, this extension lacking 
setae even at tip. 

Female. Forewing 6.0-6.5 mm long (8n). Exteriorly as described for male. Genitalia 
(Fig. 3) (3n): Posterior margin of seventh sternum inflected, overlapping ostium bursae, 
posterior corners not elaborated; eighth tergum lacking scales; middle % of ductus bursae 
sclerotized, the sclerotization encircling ductus but appearing flat and twisted, ductus 
seminalis originating just anterior to sclerotization; corpus bursae largely spinulose, with 
two equal-sized finlike signa. 


VOLUME 41, NUMBER 3 153 


Types. Holotype: female, 28 June 86, 4 km S Loja, Ecuador, reared from Juglans 
neotropica, A. Samaniego, Col. (Fig. 1), genit. prep. WEM 2710861, in U.S. National 
Museum of Natural History (USNM). Paratypes: Four males, same data as holotype except 
genit. preps. WEM 2110862 (Fig. 2), 2710862, 2738610, 2710864, and two females, same 
data as holotype except genit. preps. WEM 211086 (Fig. 3), 2810861; in USNM and 
University of Minnesota, St. Paul. 


Discussion. Gretchena garai is distinct from all known congeners 
because of the green scaling on thorax and forewings, and the absence 
of setae on the anal extensions of male cuculli. I examined three ad- 
ditional males with the same data as the types but excluded them from 
the type series because of poor condition. All specimens had some or 
many forewing scales missing; I could not tell whether they had fore- 
wing scale tufts, a generic trait. It seems likely that specimens with all 
scaling intact would appear greener. The species is named in honor of 
Robert I. Gara and his forest entomological work in Ecuador. 


ACKNOWLEDGMENTS 


I thank R. I. Gara of the U.S. Agency for International Development and the University 
of Washington, Seattle, for information and specimens; J. A. Powell, University of Cal- 
ifornia, Berkeley, for a manuscript copy of the Neotropical check list of Tortricidae 
(Powell & Razowski in press); R. L. Brown, J. B. Heppner, and two unnamed referees 
for reviewing the manuscript; and R. K. Robbins for serving as Journal editor for this 


paper. 
LITERATURE CITED 


BLANCHARD, A. & E. C. KNUDSON. 1983. New North American species of Eucosmini 
(Lepidoptera; Tortricidae). Proc. Entomol. Soc. Wash. 85:845-852. 

BRowN, R. L. 1982. Notes on Gretchena: A new species and the synonymy of Gwen- 
dolina (Lepidoptera: Tortricidae). Proc. Entomol. Soc. Wash. 84:594-602. 

CHUDNOFF, M. 1984. Tropical timbers of the world. U.S. Dept. Agr., Agr. Handbook 
607. 464 pp. 

MILLER, W. E. 1987. Guide to the olethreutine moths of Midland North America 
(Tortricidae). U.S. Dept. Agr., Agr. Handbook 660. 104 pp. 

NAUGHTON, G. G. 1970. Black walnut deformed by shoot moth. J. For. 68:28-29. 

POWELL, J. A. 1983. Tortricidae, pp. 31-41. In Hodges, R. W. (ed.), Check list of the 
Lepidoptera of America north of Mexico. E. W. Classey Ltd. & Wedge Entomological 
Research Foundation, London. 284 pp. 

POWELL, J. A. & J. RAZOwSKI. Tortricidae: Olethreutinae. In Heppner, J. B. (ed.), Atlas 
of Neotropical Lepidoptera. Checklist: Part 2. Pyraloidea—Tortricoidea. E. J. Brill, 
Leiden. In press. 


Received for publication 17 March 1987; accepted 1 May 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 154-158 


INJURY AND BIOLOGY OF THE CLEARWING BORER 
SYNANTHEDON KATHYAE ON HOLLY 


G. M. GHIDIU 


Vegetable Entomologist, Rutgers University, 
R.D. #5, Box 232, Bridgeton, New Jersey 08302 


L. VASVARY 


Extension Specialist, Ornamental Entomology, 
Rutgers University, New Brunswick, New Jersey 08903 


T. D. EICHLIN 


Insect-Taxonomy Laboratory, Division of Plant Industry, 
California Department of Food and Agriculture, Sacramento, California 95814 


AND 


J. D. SOLOMON ~ 


Southern Hardwoods Laboratory, U.S. Forest Service, 
Stoneville, Mississippi 38776 


ABSTRACT. Synanthedon kathyae Duckworth & Eichlin was reared from infested 
holly cultivars (Ilex spp.) at Bridgeton, New Jersey. This is the first reported host for S. 
kathyae. Signs of infestation were wilting, yellowing, and dying foliage. Frass galleries 
around the root collar and cracked, loose bark at the base of the plant provided further 
evidence of infestation and injury. There appears to be one generation annually. 


Additional key words: Sesiidae, Ilex spp., galleries. 


When originally described, Synanthedon kathyae Duckworth & Ei- 
chlin was known only from six specimens collected from Lewisboro 
and Long Island, New York; Oconee Co., South Carolina; and Halifax, 
Nova Scotia (Duckworth & Eichlin 1977). Subsequently, one moth was 
captured in a pheromone-baited (Z,Z isomer of 3,13-octadecadien-1-ol 
acetate) trap in Kent Co., Maryland (Neal & Eichlin 1988) and three 
males were trapped with the same attractant in Barnstable, Massachu- 
setts. Nothing was known of its hosts or biology. We report rearing this 
species from ornamental holly trees commercially grown in Cumber- 
land Co., New Jersey, describe its injury, and outline its biology. 


MATERIALS AND METHODS 


This insect first came to our notice in 1981 when we received two 
mature or nearly mature larvae collected from an American holly (Ilex 
opaca Ait.) tree on 18 October 1981 in Millville, New Jersey. The larvae 
were identified as Synanthedon, but species determination had to await 
association with characters of the adult. 

In fall 1982, a borer infestation in container-grown holly trees (Ilex 
spp.), in greenhouses at Millville was called to our attention. In No- 


VOLUME 41, NUMBER 3 155 


vember 1982, we began rearing the larval borers to the adult stage for 
identification, and recording injury and effects on the trees. Sixteen 
container-grown, four-year-old hollies (all of “Blue Angel” variety), 
including eight infested trees showing decline symptoms and eight 
apparently healthy trees, were selected from the Millville tree farm for 
study. These trees were moved to the Rutgers Research & Development 
Center at Bridgeton, New Jersey, for close and frequent observation. 

Four infested and four healthy trees were placed on wooden plat- 
forms under shaded screen emergence cages in a heated greenhouse, 
and the other eight were placed similarly in the field for natural seasonal 
emergence. The cages were constructed of wood frames and covered 
with 18 x 16-mesh galvanized screen wire 0.028 cm diam. to enclose 
an area of about 0.5 m°. The caged plants were inspected weekly until 
the first moth appeared, and daily afterward. After moths emerged, 
the infested piants were dissected to study burrowing habits and gallery 
dimensions. Adult voucher specimens were deposited in the U.S. Na- 
tional Museum of Natural History, Washington, D.C., and in the col- 
lection at the Entomology Department, Cook College, Rutgers—The 
State University, New Brunswick, New Jersey. 


RESULTS AND DISCUSSION 

The adult of S. kathyae is a bluish black moth with clear wings, 
yellow body markings, and prominent yellow bands on abdominal seg- 
ments four and five; a detailed description is presented in Duckworth 
and Eichlin (1977). The mature larva is creamy white with a dark 
brown head and light brown spiracles, and ranges from 15 to 21 mm 
in length. The egg is small, brownish, and oval. 

Initial signs of infestation were wilting and drooping of tender ter- 
minal and branch shoots. The foliage first became chlorotic to yellowish, 
and finally brown and curled. Girdled branches sometimes dropped 
their leaves (Fig. 1), while the rest of the plant remained green. Heavily 
infested plants exhibited progressive dieback with one or more dying 
limbs, and eventually the entire plant succumbed. Dieback and mor- 
tality were most noticeable in early to mid-November. 

Light brown frass was ejected from bark entrances just above the 
soil line. The frass gradually became coarse granular and accumulated 
in piles on the ground around the root collar. Raking away frass revealed 
cracked, loose bark that was easily removed to expose larvae and their 
tunnels (Fig. 2). Multiple galleries in the wood were common; up to 
six with pupal exits were observed on single plants 3 to 4 cm in diam. 
at the root collar (Fig. 3). Plants of this size infested by three or more 
larvae usually died. Galleries were irregular in shape but oval in cross- 
section, measuring 4 to 8 mm wide and 5 to 8 cm long. Most galleries 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 











Fics. 1-4. Injury and signs of Synanthedon kathyae. 1, Dieback on potted holly; 2, 


Galleries at base of main stem; 3, Pupal exit holes and galleries; 4, Pupal skin protruding 
from exit hole. 


extended 1 to 2 cm below the soil line and 3 to 7 cm above it. Galleries 
were usually kept open and clean; few contained loose frass. 

Ten male and three female moths emerged between 7 and 17 Feb- 
ruary 1983 from the four infested plants in the heated greenhouse. Two 
males and one female emerged 30-31 May 1983 from the infested 
plants in outdoor cages. Single moths of S. kathyae have been captured 
on 25 and 30 June and 14 July in Massachusetts (not previously re- 
ported); 17 and 24 July in New York; 25 June in South Carolina; and 
2 July in Maryland (Duckworth & Ejichlin 1972). Field emergence dates 


VOLUME 41, NUMBER 3 L157 


reported here are similar to those (May-July) for other clearwing species 
in the region (Neal 1982, Schread 1965, Wallace 1945, Potter & Tim- 
mons 1983, Gentry et al. 1978). 

The mature larva prepared for adult emergence by cutting a round 
exit hole 4 mm in diam., leaving only a thin bark flap as a cap. Pupation 
occurred head-upward in the gallery. Pupal exuviae protruding from 
exit holes were easily visible around the root collar and lower branches 
during the emergence season, and provided additional evidence of 
infestation (Fig. 4). Galleries, larvae of uniform size. and emergence 
within a single year suggests one generation per year. 

Several varieties of holly were attacked. Records and observations at 
the holly farm suggest that variety “Blue Angel’ (Ilex x meservae) 
was most susceptible, followed by “Nellie Stevens” (I. cornuta [Lindley] 
x I. aquifolium L.) and “Inkberry” (I. glabra L.). The varieties “Blue 
Prince” (I. aquifolium x I. rugosa [Schmidt]) and “Blue Princess” (I. 
rugosa x I. aquifolium) were less attacked. Stressed and weakened 
plants appeared more susceptible to borer attack than did healthy, 
vigorous plants. 

A question arises as to where the nursery infestation originated. The 
plants were started in the nursery at Millville and grown in pots to age 
4 years for marketing as either 30-38 cm or 38-48 cm tall plants. The 
nursery was surrounded by a mixed stand of American holly, oaks, 
white pine, and an undergrowth of shrubs. The insects might have 
moved from firewood cut in nearby stands and hauled to the nursery 
for heating, or moths might have flown from natural infestations in the 
surrounding woodland. 

The nurseryman stated that about 30 percent of the “Blue Angel” 
variety were infested during 1981-82, amounting to an estimated loss 
of $6,000. 


ACKNOWLEDGMENTS 


We thank the American Holly Farm, Millville, New Jersey; J. Johnson, Cumberland 
County Agricultural Agent; D. DeBlois, New Jersey Department of Agriculture; and 
Rutgers Research & Development Center farm manager S. B. Sheppard for providing 
facilities and technical assistance; also D. Weisman, National Museum of Natural History, 
for identifying larvae. New Jersey Agricultural Experiment Station Publication No. D-08120- 
03-87 supported by State Funds and by the United States Hatch Act. 


LITERATURE CITED 


DucKworTH, W. D. & T. D. EICHLIN. 1977. Two new species of clearing moths 
(Lepidoptera: Sesiidae) from eastern North America clarified by sex pheromones. J. 
Lepid. Soc. 31:191-196. ; 

Genny, C. GC. RL. HoLrLtoway & D. K. PoLLeT. 1978. Pheromone monitoring of 
peach tree borers and lesser peach tree borers in South Carolina. J. Econ. Entomol. 
71:247-258. 


158 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NEAL, J. W., JR. 1982. Rhododendron borer: A worthy competitor. J. Am. Rhododendron 
Soc. 36:57-61. 

NEAL, J. W., JR. & T. D. EICHLIN. 1983. Seasonal response of six male Sesiidae of woody 
ornamentals to clearwing borer (Lepidoptera: Sesiidae) lure. Environ. Entomol. 12: 
206-209. 

Potter, D. A. & G. M. Timmons. 1988. Flight phenology of the dogwood borer 
(Lepidoptera: Sesiidae) and implications for control in Cornus florida L. J. Econ. 
Entomol. 76:1069-1074. 

SCHREAD, J. C. 1965. Dogwood borer. Conn. Agr. Exp. Stn. Cir. 199. 4 pp. 

WALLACE, P. P. 1945. Biology and control of the dogwood borer, Thamnosphecia 
scitula Harris. Conn. Agr. Exp. Stn. Bull. 488:373-395. 


Received for publication 19 March 1987; accepted 10 June 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 159-165 


THE ROLE OF NECTAR SOURCE DISTRIBUTION IN 
HABITAT USE AND OVIPOSITION BY THE 
TIGER SWALLOWTAIL BUTTERFLY 


DAVID W. GROSSMUELLER AND ROBERT C. LEDERHOUSE! 


Department of Zoology and Physiology, Rutgers University, 
Newark, New Jersey 07102 


ABSTRACT. In northern New Jersey, larval hosts of the polyphagous butterfly Papilio 
glaucus L. are widely distributed but nectar plants are clumped. Females were rarely 
captured in an area rich in preferred hosts during the spring generation, when the area 
lacked concentrations of nectar plants. During summer, second generation females fed 
and oviposited in the study area while thistles bloomed. The locations of hosts receiving 
eggs correlated significantly with locations of nectar plants. Such an ovipositional pattern, 
although probably not maladaptive for larvae, appears to be unrelated to improved larval 
growth or survival. 


Additional key words: Papilionidae, Papilio glaucus, adult resources, activity centers. 


The distribution of individuals of a butterfly population generally 
reflects the distribution of their resources. Sources of adult or larval 
nutrition, or areas facilitating mating or roosting, may all occur in the 
same habitat as in the Jasper Ridge colonies of Euphydryas editha 
(Ehrlich 1965). In this case the population was local and sedentary 
(Gilbert & Singer 1973). More commonly, different requirements are 
met in different habitats. Other populations of E. editha were much 
less localized due to the dispersed distribution of nectar sources (Gilbert 
& Singer 1973) or local defoliation of larval host plants (Murphy & 
White 1984). In central Sweden, Leptidea sinapis females regularly 
moved from a woodland nectar foraging habitat to a meadow ovipo- 
sition habitat (Wiklund 1977). Some habitats that appear to be rich in 
larval resources may be rarely used (Singer 1971, Cromartie 1975, Chew 
1977, Rausher 1979). Areas of otherwise suitable habitat may not be 
used if they lack a limited resource such as adult nectar plants (Murphy 
1983). 

In any one habitat, butterflies often do not oviposit on all of their 
potential larval hosts (Singer 1971, Wiklund 1975, Smiley 1978), nor 
use all of the habitats in which preferred hosts grow (Singer 1972, 
Cromartie 1975). Selective oviposition has been interpreted to favor 
better larval growth or survival (Ehrlich & Raven 1964, Wiklund 1975, 
Holdren & Ehrlich 1982). Although this has been clearly demonstrated 
in some cases (Rausher 1980, Rausher & Papaj 1983, Williams 1983), 
in other cases, no advantage for the larvae has been shown (Chew 1977, 
Rausher 1979, Courtney 1981, 1982). Certain hosts may favor high 


1 Present address: Department of Entomology, Michigan State University, East Lansing, Michigan 48824 


160 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


oviposition rates, and higher larval mortality is offset by the greater 
number of eggs (Rausher 1979, Courtney 1982). If hosts are widely 
distributed and of similar overall quality, selective oviposition may 
represent the response of females to other factors such as restricted 
adult resources (Murphy et al. 1984). This suggests that in many cases 
the distribution of eggs is a compromise between larval and adult needs. 

The tiger swallowtail, Papilio glaucus L., is the most polyphagous of 
the over 500 species of Papilionidae (Scriber 1973), using hosts from 
over 14 plant families (Tietz 1972). Northern New Jersey is near the 
limit of bivoltine populations of P. glaucus (Hagen & Lederhouse 1985), 
and in this area, P. glaucus females show a strong preference for tu- 
liptree, Liriodendron tulipifera L. (Grossmueller & Lederhouse 1987), 
although several other suitable hosts are abundant (Grossmueller 1984). 
In northern New Jersey, they also oviposit on the W and S sides of trees 
within 3 m of the ground (Grossmueller & Lederhouse 1985). Such 
selective oviposition greatly increases the probability of completing two 
generations during normal and cooler than normal years. Potential 
larval hosts are abundant in this area, and P. glaucus populations never 
reach densities high enough to deplete their larval food supply. Our 
question in the present report was whether the temporal and spatial 
distribution of adult resources affect which habitats and individual hosts 
are selected for oviposition. 


MATERIALS AND METHODS 

The study area was located in Flanders, Morris Co., New Jersey, and 
consisted of a 5300 m? extension of a larger field (Grossmueller & 
Lederhouse 1985). It was surrounded on three sides by woods, and had 
progressed to late old-field succession. Potential hosts such as tuliptree, 
wild black cherry (Prunus serotina Ehrh.), and white ash (Fraxinus 
americana 1.) were abundantly scattered throughout the open field, 
around the edges and in the surrounding woods. The only major source 
of nectar in or around the study site was bull thistle, Cirsium vulgare 
L., which was in bloom from mid-July through August. The locations 
of all flowering bull thistles and of all potential hosts within the study 
site were mapped. The site was divided into 23 plots (10 m x 20 m) 
to facilitate relocating marked hosts (Grossmueller 1984). 

Adult P. glaucus were captured and marked from 1 May until 15 
September each year of the study. Date, time of day, atmospheric 
conditions, age, sex, and behavior at time of capture were recorded. 
No butterflies were removed from the study site. Eggs were sampled 
four times per week during June through August 1980-82. The upper 
surface of all potential host leaves at heights of 5 m or less were examined 
for each host along a 310 m long, 2 m wide transect through the study 
area. Additional sampling of taller hosts was described in Grossmueller 


VOLUME 41, NUMBER 8 161 


TABLE 1. Behavior of Papilio glaucus adults immediately before capture within the 
study area. 


Percentage 
Brood Sex n Pudding ~+~‘Filying=~~—~*«~Necttringe 
1980-1 Male 43 83.7 9.3 Le 
Female 4 == 25.0 75.0 
1980-2 Male 35 —_ 11.4 88.6 
Female 42 — Tea 92.9 
1981-1 Male Pall 85.7 9.5 4.8 
Female 1 — — 100.0 
1981-2 Male 3 100.0 — —— 
Female 6 — 16.7 83.3 


and Lederhouse (1985), but relatively few eggs were laid above 3 m. 
Because 88.2% of the eggs found within the study area were on tuliptree 
(Grossmueller 1984), only the relation between the location of tuliptrees 
on which females oviposited and the location of flowering thistles was 
analyzed. 


RESULTS 


Although the study site was selected for its abundance of potential 
P. glaucus larval hosts, only 5 eggs (4.6% of the yearly total) were found 
in it during the first generation of 1980. Tiger swallowtails were fre- 
quently encountered in the area, and 47 were captured and marked. 
However, 43 (91.5%) of these were males, nearly all of which had been 
puddling along a swimming pool at the edge of the study area (Table 
1). Puddling males form conspicuous groups on moist soil where they 
ingest sodium, amino acids, and possibly other nutrients (Arms et al. 
1974, Berger & Lederhouse 1985). 

During the second generation of 1980, 103 eggs were found within 
the study area. Tiger swallowtails were somewhat more abundant; 77 
were captured and marked. The major differences were that only 45.5% 
of the captures were males, and 90.9% of all captures were nectaring 
on bull thistle (Table 1). A significantly greater proportion of males 
captured during the second generation were nectaring (x?, P < 0.001). 
However, the distribution of captured females did not differ by behavior 
between the two broods. The number of eggs discovered each gener- 
ation was distributed differently from the total number of P. glaucus 
captures by brood (x?, 2 x 2 contingency table, P < 0.001) but did not 
differ from the distribution of female captures (P > 0.50). 

The use of the study site during the first generation of 1981 was 
similar to that brood in 1980. Of the 22 P. glaucus that were captured, 
95.5% were male, and 85.7% of the males had been puddling when 
captured. Only one egg was found during this brood. During June 1981, 


162 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the State of New Jersey sprayed the entire Flanders area with a for- 
mulation of Bacillus thuringiensis to kill gypsy moth caterpillars. Tiger 
swallowtail caterpillars are easily killed by this insecticide (Grossmueller 
unpubl. results). Only nine captures were made during the second 
generation of 1981, and only six eggs were found. However, two-thirds 
of the captures were females, which was consistent with the previous 
year. The population level of tiger swallowtails continued to be very 
low throughout 1982. No eggs were found during the first generation 
of 1982, but nine were found at the site during the second generation. 

Despite differences in overall population size, the pattern of study 
site use was similar from year to year. Although host trees were of 
similar availability and quality throughout oviposition periods, females 
were attracted to and oviposited within the study area mainly when 
thistles were in bloom. Males were attracted to the area to puddle during 
first broods, and to feed on nectar when thistles were blooming. 

Since blooming thistles attracted females to the area during the second 
generation, the influence of thistle location on the particular host plants 
chosen for oviposition was investigated. Using the large sample from 
the second brood of 1980, numbers of eggs, flowering thistles, and 
tuliptree hosts were determined for each of the 23 plots. There was no 
relation between number of thistles and number of tuliptrees in the 
plots (n = 28, r = 0.12, P > 0.50). Surprisingly, number of eggs was 
not correlated significantly with number of host trees (r = 0.11, P > 
0.50). Since eggs must be on a host tree, we excluded plots without 
hosts from analysis of the relation between eggs and thistles. The number 
of eggs was correlated significantly with number of thistles for plots 
with at least one host (n = 17, r = 0.55, P < 0.05). 

A modified nearest-neighbor analysis was used to compare the mean 
distance of each egg (n = 108) to the nearest thistle. The nearest- 
neighbor value was 0.27, which indicates a significant clumped spatial 
association between eggs and thistles. The value for all host trees was 
0.76, which indicates a random distribution. 


DISCUSSION 


In the area surrounding the study site, tiger swallowtails are probably 
not limited by availability of suitable larval hosts. In northern New 
Jersey, many previously cultivated fields have undergone succession as 
residential communities have largely replaced agriculture. Many of the 
host species are early colonizers of abandoned fields, resulting in young 
stands of tuliptree or white ash. Even at high population levels, P. 
glaucus uses only a small portion of available hosts. For example, in 
the second brood of 1980, there were 108 eggs oviposited on tuliptrees 
within the study area. However, there were approximately 50,800 tu- 


VOLUME 41, NUMBER 3 163 


liptree leaves that were suitable in condition and location for larval 
development (Grossmueller 1984). Thus, females can afford to ignore 
entire areas of host plants in favor of areas containing nectar sources. 

In contrast, adult nutrition sources are more restricted. Male tiger 
swallowtails often form large aggregations at puddling sites (Arms et 
al. 1974). A puddling site was the prime focus of P. glaucus activity 
during the first broods at our study site. In a study of P. glaucus behavior 
in upstate New York, Berger (1986) found much higher frequencies of 
occurrence near nectar sources than at locations with abundant host 
plants but devoid of nectar plants. Our impression is that most of the 
naturalized nectar plants used during both generations in New Jersey 
tend to be clumped. Native hosts such as basswood (Tilia americana 
L.) and various briars (Rubus spp.) are similarly clumped. When larval 
hosts are widely distributed and adult resources are localized, female 
activity and oviposition are greatest near the adult resources (Murphy 
1983, Murphy et al. 1984). 

Tiger swallowtails are capable of considerable movement in a short 
time (Lederhouse 1982a). Movements in excess of 5 km between cap- 
tures are not uncommon for marked males, with one detected displace- 
ment of over 2 km in | h (Lederhouse unpubl. results). Although females 
could move from host patches to nectar patches and back again, they 
appear not to move as much as males (Lederhouse 1982a). Since many 
nectar plant concentrations may be included in suitable ovipositional 
habitat, such movement may be unnecessary. 

Oviposition in areas of increased male activity is often reduced due 
to male harassment (Shapiro 1970, Lederhouse 1982b). In Euphydryas 
chalcedona, courtship by nectar-feeding males may reduce the number 
of eggs laid in the immediate vicinity of nectar plants (Williams 1983). 
However, disturbed females move only short distances to resume ovi- 
position, producing the overall relation of egg location to nectar plant 
location (Murphy et al. 1984). Whether this pattern is detected (Murphy 
et al. 1984) or not (Williams 1983) is a function of what scale is used. 
When harassment does occur, females may balance problems of inter- 
ference against costs of extra flight to and away from nectar plants. 

In our study, the number of males within the study area was similar 
for the two generations of 1980. Therefore, lack of females and eggs 
at the site during the first brood was probably not due to avoidance of 
males. Furthermore, the presence of abundant nectar plants during the 
second generation attracted females to the area. The difference in 
number of females within the study area between the two generations 
largely accounted for the difference in number of eggs discovered. 
During the second generation, proximity to a thistle or thistle patch 
increased the probability that a preferred host would have eggs on it. 


164 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Thus, nectar availability controls the local distribution of tiger swal- 
lowtail larvae in northern New Jersey. 

Results from this study and others (Rausher 1979, Courtney 1982, 
Murphy 1983, Murphy et al. 1984) support the contention that the 
distribution of eggs in many butterfly species is a compromise between 
larval and adult needs. The relative importance of various components 
in this trade-off will determine the nature of the observed ovipositional 
pattern. 


ACKNOWLEDGMENTS 


We thank T. Berger, S. Codella, R. Hagen, and M. Rausher for helpful comments on 
drafts of this manuscript. 


LITERATURE CITED 


ARMS, K., P. FEENY & R. C. LEDERHOUSE. 1974. Sodium: Stimulus for puddling behavior 
by tiger swallowtail butterflies, Papilio glaucus. Science 185:372-374. 

BERGER, T. A. 1986. Habitat use and reproductive ecology of the eastern tiger swal- 
lowtail, Papilio glaucus L. Dissertation, Rutgers University, Newark, New Jersey, 
USA. 109 pp. Diss. Abstr. Int. microfilm order No. 8616563. 

BERGER, T. A. & R. C. LEDERHOUSE. 1985. Puddling by single male and female tiger 
swallowtails, Papilio glaucus L. (Papilionidae). J. Lepid. Soc. 39:339-340. 

CHEW, F. S. 1977. Coevolution of pierid butterflies and their cruciferous foodplants. 
II. The distribution of eggs on potential foodplants. Evolution 31:568-579. 

CouRTNEY, S. P. 1981. Coevolution of pierid butterflies and their cruciferous foodplants. 
III. Anthocharis cardamines (L.) survival, development and oviposition on different 
hostplants. Oecologia (Berl.) 51:91-96. 

1982. Coevolution of pierid butterflies and their cruciferous foodplants. IV. 
Crucifer apparency and Anthocharis cardamines (L.) oviposition. Oecologia (Berl.) 
02:258-265. 

CROMARTIE, W. J. 1975. The effect of stand size and vegetational background on the 
colonization of cruciferous plants by herbivorous insects. J. Appl. Ecol. 12:517-533. 

EHRLICH, P. R. 1965. The population biology of the butterfly, Euphydryas editha. I. 
The structure of the Jasper Ridge colony. Evolution 19:327-336. 

EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: A study in coevolution. 
Evolution 18:586-608. 

GILBERT, L. E. & M. C. SINGER. 1973. Dispersal and gene flow in a butterfly species. 
Am. Nat. 107:58-78. 

GROSSMUELLER, D. W. 1984. Factors affecting voltinism in the tiger swallowtail Papilio 
glaucus. Dissertation, Rutgers University, Newark, New Jersey, USA. 114 pp. Diss. 
Abstr. Int. microfilm order No. 8422557. 

GROSSMUELLER, D. W. & R. C. LEDERHOUSE. 1985. Oviposition site selection: An aid 
to rapid growth and development in the tiger swallowtail butterfly, Papilio glaucus. 
Oecologia (Berl.) 66:68-73. 

1987. Host selection in the tiger swallowtail butterfly, Papilio glaucus, related 
to its potential to complete an additional brood. Ecol. Entomol. Submitted. 

HAGEN, R. H. & R. C. LEDERHOUSE. 1985. Polymodal emergence of the tiger swallow- 
tail, Papilio glaucus (Lepidoptera: Papilionidae): Source of a false second generation 
in central New York State. Ecol. Entomol. 10:19-28. 

HOLDREN, C. E. & P. R. EHRLICH. 1982. Ecological determinants of food plant choice 
in ie checkerspot butterfly Euphydryas editha in Colorado. Oecologia (Berl.) 52: 
417-423. 

LEDERHOUSE, R. C. 1982a. Factors affecting equal catchability in two swallowtail 
butterflies, Papilio polyxenes and P. glaucus. Ecol. Entomol. 7:379-383. 








VOLUME 41, NUMBER 3 165 





1982b. Territorial defense and lek behavior of the black swallowtail butterfly, 
Papilio polyxenes. Behav. Ecol. Sociobiol. 10:109-118. 

Murpnuy, D. D. 1983. Nectar sources as constraints on the distribution of egg masses 
by the checkerspot butterfly, Euphydryas chalcedona (Lepidoptera: Nymphalidae). 
Environ. Entomol. 12:463—466. 

Murpny, D. D., M. S. MENNINGER & P. R. EHRLICH. 1984. Nectar source distribution 
as a determinant of oviposition host species in Euphydryas chalcedona. Oecologia 
(Berl.) 62:269-271. 

Murpny, D. D. & R. R. WuHiTe. 1984. Rainfall, resources, and dispersal in southern 
populations of Euphydryas editha (Lepidoptera: Nymphalidae). Pan-Pac. Entomol. 
60:350-354. 

RAUSHER, M. D. 1979. Larval habitat suitability and oviposition preference in three 
related butterflies. Ecology 60:503-511. 

1980. Host abundance, juvenile survival and oviposition preferences in Battus 
philenor. Evolution 34:342-355. 

RAUSHER, M. D. & D. R. Papaj. 1983. Demographic consequences of discrimination 
among conspecific host plants by Battus philenor butterflies. Ecology 64:1402-1410. 

SCRIBER, J. M. 1973. Latitudinal gradients in larval specialization of the world Papil- 
ionidae (Lepidoptera). Psyche 80:355-3738. 

SHAPIRO, A. M. 1970. The role of sexual behavior in density-related dispersal of pierid 
butterflies. Am. Nat. 104:367-372. 

SINGER, M. C. 1971. Evolution of food-plant preference in the butterfly Euphydryas 
editha. Evolution 25:383-389. 

1972. Complex components of habitat suitability within a butterfly colony. 
Science 176:75-77. 

SMILEY, J. 1978. Plant chemistry and the evolution of host specificity: New evidence 
from Heliconius and Passiflora. Science 201:745-747. 

Tietz, H. M. 1972. An index to the described life histories, early stages, and hosts of 
the Macrolepidoptera of the continental United States and Canada. 2 vols. Allyn, 
Sarasota, Florida. 1041 pp. 

WIKLUND, C. 1975. The evolutionary relationship between adult oviposition preferences 
and larval host plant range in Papilio machaon L. Oecologia (Berl.) 18:185-197. 
1977. Oviposition, feeding and spatial separation of breeding and foraging 

habitats in a population of Leptidea sinapis (Lepidoptera). Oikos 28:56-68. 

WILLIAMS, K. S. 1983. The coevolution of Euphydryas chalcedona butterflies and their 
larval host plants III. Oviposition behavior and host plant quality. Oecologia (Berl.) 
06:336-340. 











Received for publication 11 May 1987; accepted 18 June 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 166-167 


GENERAL NOTE 


PENSTEMON DIGITALIS (SCROPHULARIACEAE), A NEW 
FOOD PLANT RECORD FOR HAPLOA CONFUSA (ARCTIIDAE) 


Additional key words: alkaloid, boschniakine. 


The confused Haploa moth (or Lyman’s Haploa, Haploa confusa Lyman) is common 
during July in tallgrass prairie habitats in E-central Illinois. However, little information 
is available on its larval food plants. Haploa confusa is not listed by Tietz (1972, An index 
to the described life histories, early stages, and hosts of the Macrolepidoptera of the 
continental United States and Canada, A. C. Allyn, Sarasota, Florida, 536 pp.), and Holland 
(1968, The moth book, Dover Publications, New York, 479 pp.) makes no mention of 
food plants. Covell (1984, A field guide to the moths of eastern North America, Houghton 
Mifflin, Boston, 496 pp.) lists only “hound’s-tongue”’ as a food plant. Several herbaceous 
plant species are commonly called hound’s-tongue, but the name best applies to Cyno- 
glossum officinale L. (Boraginaceae), a herbaceous plant of European origin widely 
naturalized in the United States (Lyons, 1901, Plant names scientific and popular, Nelson, 
Baker and Co., Detroit, 630 pp.). Other Haploa species feed on a variety of plants, 
including Eupatorium (Compositae), Salix (Salicaceae), and Triosteum (Caprifoliaceae) 
species (Holland, above). 

Foxglove penstemon (Penstemon digitalis Nutt., Scrophulariaceae) is a common bien- 
nial forb in tallgrass prairie habitats of the Ecological Research Area maintained near 
Urbana, Illinois, by the University of Illinois. Other dominant plant species in the habitat 
include big bluestem (Andropogon gerardii (Vitm.), Gramineae), Indian grass (Sorghas- 
trum nutans (L.), Gramineae), and lespedeza (Lespedeza cuneata (Dum.-Cours.) G. Don, 
Leguminosae) (Lindroth & Batzli, 1984, J. Mamm. 65:600-606). On 1 May 1984, while 
collecting P. digitalis for analysis of alkaloid constituents, I found a second or third stage 
larva resting on a P. digitalis rosette leaf. The leaf was partially eaten and some larval 
frass adhered to it. The larva was taken into the laboratory and reared through its 
remaining stadia on leaves of P. digitalis. The larva was maintained in a glass Petri dish 
and fed clumps of rosette leaves until the final stadium, at which time I transferred it to 
a small Plexiglas cage covering a potted P. digitalis plant. The larva exhibited an unusual 
feeding behavior; it started feeding at the tip of a leaf, then cut a narrow (<1 cm) swath 
down the center of the leaf, consuming the midrib and a narrow band of tissue on both 
sides. After pupation, the insect was not visible on the plant or soil surface. Two to three 
weeks after pupation, a female H. confusa emerged. 

Use of P. digitalis by H. confusa is interesting because the plant contains especially 
high levels of boschniakine (a pyridine monoterpene alkaloid) during the larval feeding 
period (Lindroth et al., 1986, Biochem. Syst. Ecol. 6:597-602). Alkaloids commonly occur 
in the food plants of Haploa species, including cynoglossophine, heliotrine, lasiocarpine, 
and platyphilline in Cynoglossum officinale, and echinatine and trachelanthamidine in 
Eupatorium maculatum L. (Willaman & Li, 1970, Lloydia 33, Suppl. No. 3A:1-286). In 
addition to alkaloids, Penstemon species commonly contain a variety of iridoid glycosides, 
such as ajugol and catalpol (Junior, 1983, Planta Medica 47:67-70), although it is not 
known whether P. digitalis contains the compounds. Both alkaloids and iridoid glycosides 
are deterrent or toxic to many insects (Robinson, 1979, pp. 413-448 in Rosenthal & 
Janzen, Herbivores: Their interaction with secondary plant metabolites, Academic Press, 
New York, 718 pp.; Bernays & De Luca, 1981, Experientia 37:1289-1290). Thus H. 
confusa larvae probably have physiological or biochemical adaptations that enable them 
to avoid the effects of these potentially toxic compounds. Alkaloids and iridoid glycosides 
are sequestered as defensive compounds in other lepidopteran species (Duffey, 1980, Ann. 
Rey. Entomol. 25:447-477; Bowers, 1980, Evolution 34:586-600). The apparent apose- 
matic coloration of H. confusa larvae, black with orange longitudinal stripes, indicates 
that they may sequester alkaloids or iridoid glycosides from P. digitalis. 


VOLUME 41, NUMBER 3 167 


I thank G. L. Godfrey (Illinois Natural History Survey, Champaign) for identifying 
the moth; W. C. Capman generated enthusiasm and provided advice for rearing the 
larva. 


RICHARD L. LINDROTH, Department of Entomology, University of Wisconsin, 1630 
Linden Drive, Madison, Wisconsin 53706. 


Received for publication 22 January 1987; accepted 13 May 1987. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 168 


BOOK REVIEWS 


MEXICAN LEPIDOPTERA: EURYTELINAE I, by R. de la Maza E. and R. Turrent D., color 
photographs by R. Doniz. 1985. iii + 44 pp., 43 maps, 19 plates (8 in color), 4 unnumbered 
halftone photos in text. In English. Sociedad Mexicana de Lepidopterologia, AC Publi- 
caciones Especiales 4. Distributed exclusively in United States by Entomological Reprint 
Specialists, P.O. Box 77224, Dockweiler Station, Los Angeles, California 90007. Paper- 
bound, $25. 


Not since the Biologia Centrali Americana and Seitz’ Macrolepidoptera of the World 
has Mexico’s diverse lepidopteran fauna been reviewed. The present work is the first in 
a proposed series to rectify that situation, the authors’ intent “... to make the Mexican 
Lepidoptera known internationally.” They chose, for their initial volume, the subfamily 
Eurytelinae (family Nymphalidae), a group about which a number of generic revisions 
and life history studies covering Mexican species have appeared during the past 40 years. 

There are good points to the work. A male genitalic drawing and at least one color 
photograph illustrates each of the 44 species (Perisama mexicana Hoffmann, known from 
but a single specimen, receives only a halftone photo in the text section). There are no 
female genitalic drawings. Seven new subspecies are described and illustrated with half- 
tone photographs. The new forms are based on minor differences in coloration, though 
genitalic characters are sometimes mentioned but not elaborated. The diagnoses fulfill 
the Rules of Zoological Nomenclature but I should like to see a supplemental paper 
presenting more detailed comparisons with related subspecies. Particularly valuable are 
species keys, drawings of wing nomenclature and venation (Plate II), and diagnostic 
drawings of hard-to-differentiate species in Diaethria and Myscelia (Plate I). Also, a 
distribution map is given for every species. Part IV admirably summarizes recent literature 
on known food plants and immature stages. Because such data are available for only 14 
species in 12 of 16 genera covered, the authors indirectly point to the need for additional 
research on immature stages. 

Unfortunately, annoying problems seriously reduce the work’s value. I emphasize, 
however, that most of these involve presentation, not content. Abundant typographical 
and grammatical errors make the reading almost painful. For example, on p. 33 there 
appears, “... drinking in wet rocks and mudd, and resting on shaddy roks of the creek 
wall ....”° I cannot determine whether the authors or a third party wrote the English 
version (more than once, the Spanish y appears instead of and). At any rate, for Ento- 
mological Reprint Specialists to have permitted a presentation so ineptly edited to go to 
press is deplorable. 

I appreciate the maps, but they lack localities and outlines of the Mexican States. It 
appears that three base templates, none with northern or southern borders of the country 
shown, were used for the maps. Future workers with the Mexican fauna will still have 
to examine specimen labels and search sparse and scattered literature. 

Although the halftone photos are excellent, the color photographs, almost without 
exception, have shadows (especially dark on Plate XIX). Comparisons with Moths of 
North America (MONA) are inevitable, but the quality of the present work does not 
approach MONA’s standards. 

Finally, no tribal and generic keys nor synonymies at any taxonomic level are provided. 
The species keys are clear and easy to use, and perusal of the photographs suffice for 
species identification. However, the work could have advanced the art and science of 
euryteline systematics had it carefully outlined supraspecific group differences beyond 
a brief, noncomparative summaries provided. Omission of synonymies seems inexcus- 
abDie. 

In summary, this first offering in the series goes far toward filling an important gap in 
our knowledge of world Lepidoptera. As such, the work must take its place beside the 
few omnibus reviews now available. If only care had been taken to imbue it with more 
production quality. 


SANFORD R, LEFFLER, 4701 15th Ave. NE, #6, Seattle, Washington 98105. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 169 


NOCTUELLES ET GEOMETRES D’EUROPE, Volume II, by Jules Culot. Republication by 
Apollo Books, Lundbyvej 36, DK-5700-Svendborg, Denmark. 243 pp., 81 color plates. 
DKK 1,380.- for Vols. I-II, and also for Vols. I-IV. DKK 2,500.- for all four. 


This book and its companion volumes are a complete facsimile of the original edition 
(1909-1920) of Culot’s work without additional text. The language is French. This par- 
ticular volume covers quadrifine noctuid subfamilies and a significant proportion of the 
trifines: Cuculliinae, Heliothinae, some Hadeninae, and some Amphipyrinae. Vol. II com- 
pletes Noctuidae, and the remaining two cover Geometridae. 

For each species covered there are diagnoses and brief details of distribution, biology, 
and phenology. All are illustrated in color with Culot’s skilled and accurate artwork. 

At the time of publication, the series was intended to give a compiete coverage of the 
two families for Europe. The author was in contact with notable lepidopterists of his 
time, particularly Charles Oberthtir, and the work was well received by contemporary 
reviewers who commented both on the crying need for such a work and on the then 
unsurpassed quality of the illustrations (Entomol. 42:326-327). It is now presented un- 
changed to a new readership 70 years after its original publication. 

I cannot comment on its value as a collector’s item. The vagaries of book collectors are 
unfathomable; the practicing entomologist can only fulminate when useful but old and 
rare books are sold at prices inflated beyond the range of his pocket. But both will consider 
the quality of the color plates, the former on faithfulness to the original, the latter on 
faithfulness to life. The former is likely to be more disappointed because the reproductions 
lack some of the vivid quality of the originals, and some, but by no means all, have 
acquired an unfortunate fine speckling. The silvery white patches on some Cucullia species 
on Plate 62 have become clouded with pale ochreous brown, perhaps because the printer 
considered the accurate originals to look “washed out’. 

Even with this loss of quality, the practicing entomologist will find that the illustrations 
stand comparison with most similar modern artwork and reproduction. However, as a 
means to identification, skilled reproduction of sharp, accurate color photography must 
take precedence. 

The nomenclature used in the work has, of course, become dated in many instances, 
particularly generic combinations. The book would thus have been much more valuable 
to practicing entomologists had the publishers commissioned a specialist on European 
Lepidoptera to write an introductory section including a modern checklist (with cross- 
reference to text and plates) of the species covered, and comments where advances in 
systematics have changed the picture (discovery of species complexes), or where further 
systematic work needs to be done. 

Now, as then, there is great need for a modern, comprehensive, authoritative, well 
illustrated treatment of the European macrolepidoptera fauna, with full investigation of 
the plethora of scattered type material. Reproduction of a 70-year-old work is no substitute, 
but it may serve to concentrate the minds of European lepidopterists on the need for a 
coordinated continent-wide campaign rather than fragmentary regional skirmishes. The 
proposed Faunistica Lepidopterorum Europaeorum (Nota Lepid. 4:90-94) is a promising 
move in this direction. 


JEREMY HoL_oway, C. A. B. International Institute of Entomology, 56 Queen's Gate, 
London SW7 5JR, United Kingdom. 


Journal of the Lepidopterists’ Society 
41(3), 1987, 170-172 


OBITUARY 


ARTHUR CHARLES SHEPPARD (1902-1987) 


The staff of the Lyman Entomological Museum and all who knew Arthur Charles 
Sheppard feel a deep sense of loss with his death on 10 March 1987. 

Mr. Sheppard was born at Henley-on-Thames, Oxford, England, on 4 August 1902. At 
the age of 5, he came with his parents to live in Montréal. His formal education in schools 
was limited; he left school at the age of 13 to obtain gainful employment. He did, however, 
continue his education at night schools to prepare himself for the business world. He was 
employed by the Canadian Import Company in Montréal from February 1920 until he 
retired in April 1968 as Chief Accountant. 

Charles Sheppard’s father provided the “spark” that initiated his life-long interest and 
avocation in the world of nature. Together they observed and studied birds on weekly 
Sunday morning walks. In those days, Montréal was much different than it is now and 
observations of birds and insects could be made by strolling along Decarie Boulevard. At 
the age of 9, Sheppard became fascinated with insects and their ways and started his first 
insect collection. 

By 1917, he began more intensive study of insects and exhibited a small collection of 
butterflies at a Y.M.C.A. hobby show, and similar exhibits at Boy Scout hobby shows. It 
was at one of these shows that he met Albert F. Winn, then Curator of the Lyman 
Collection of insects at the Redpath Museum, McGill University. Winn urged him to join 
the Montréal Branch of the Entomological Society of Ontario, which he did in June 1918. 
The meetings of the Branch provided the opportunity for the young man to meet many 
entomologists, amateur and professional. His association with these people with common 
interests was all that was needed to develop and intensify his enduring interest in the 
Lepidoptera, particularly of the eastern Canadian species. 

Sheppard was Secretary-Treasurer of the Montréal Branch of the Entomological Society 
of Ontario during 1937 and 1946-49, and Treasurer in 1950. Following revival of the 
Entomological Society of Québec, he was Treasurer in 1951 and then Treasurer of the 
Montréal Branch from 1952 to 1954. At the time of his death, he had been a member 
of the Society continuously for more than 68 years. The Québec Society honored him by 
conferring on him the status of Honorary Member. 

He was a charter member of the Lepidopterists’ Society and a member of the Québec 
Society for the Protection of Birds, since 1935. From 1938 to 1947, he was a member of 
the Southern California Academy of Sciences. 

His association with the Lyman collection of insects began in 1918 and continued over 
the years as many Entomological Society meetings were held at the Redpath Museum in 
the “Lyman Room’. He was closely associated with George A. Moore, Curator of the 
Lyman collections after the death of Winn in 1935. 

The Lyman collection of insects was moved to Macdonald College in 1961, to become 
the Lyman Entomological Museum, when I was appointed Curator. 

Sheppard’s association with the collection continued, and following his retirement from 
the Canadian Import Company, he was appointed Honorary Curator (Lepidoptera), a 
position he held until his death. He was also a member of long standing on the Lyman 
Entomological Committee of McGill University, the committee which served many years 
as overseer of Museum operations. 

Sheppard’s work on Lepidoptera was invaluable. The Museum, known since 1972 as 
Lyman Entomological Museum and Research Laboratory, benefitted not only by his skill 
and patience in pinning specimens of the tiniest Microlepidoptera and in rearing speci- 
mens, but also by his arranging and curation. He donated many thousands of specimens 
to the Museum, and when the Museum purchased his personal collection (approximately 
30,000 specimens) we determined that he had already donated at least that many spec- 
imens. The result is that the Lyman Lepidoptera is the finest and most complete collection 
of Québec lepidoptera in existence. Sheppard also discovered several new species of 
insects. He described one species, and a number of other species were named in his honor. 

All of us who worked closely with Charles Sheppard in the Museum, particularly 


VOLUME 41, NUMBER 3 171 





Arthur Charles Sheppard 


Honorary Curator D. Neil Duffy and I, knew him as a shy, unassuming individual, always 
ready to share his seemingly endless store of knowledge, yet too modest to accept the 
credit that he so richly deserved. He was a true and loyal friend and in every way a real 
gentleman. He will be sorely missed by all of us. 

He is survived by his wife Dorothy, daughter Helen (Mrs. C. W. Walker), grand- 
daughters Brenda, Laurie, and Nancy, his sister Dorothy and a number of step-children 
and their families. We sympathize with them and share their loss. 


PAPERS BY A. C. SHEPPARD 


1934. A new species of the Genus Incisalia. Can. Entomol. 66:141-142. 

1945. A new record for Canada (Lepidoptera). Can. Entomol. 77:55. 

1959. Caradrina morpheus, a new record for North America of a European Moth (Noc- 
tuidae). J. Lepid. Soc. 13:77. 

1959. Mr. George A. Moore. Can. Dep. Agric., Res. Br. Entomol. Newsletter 37(10):2-3. 

1970. Palaearctic Lepidoptera new to the Province of Québec. Ann. Soc. Entomol. Québec 
15:14-16. 

1974. Palaearctic Lepidoptera in the Province of Québec. Ann. Soc. Entomol. Québec 
19:119-120. 

1975. Lygris associata Bork., a new record for North America (Lepidoptera: Geometri- 
dae). Ann. Soc. Entomol. Québec 20:7. 

1975. Sheppard, A. C. & V. R. Vickery. Types and type designations of North American 
Lepidoptera in the Lyman Entomological Museum Collection. Can. Entomol. 107: 
1129-1182. 


172 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1977. The pale form of Thymelicus lineola (Hesperidae) in the Province of Québec, 
Canada. Ann. Soc. Entomol. Québec 22:215-216. 


PAPERS IN WHICH SPECIES WERE NAMED IN HONOR OF A. C. SHEPPARD 


FREEMAN, T. N. 1944. A new psychid from Quebec (Lepidoptera: Psychidae). Can. 
Entomol. 76:186-187 (Hyaloscotes sheppardi Freeman). 

McDuNNOUGH, J. 1929. Some apparently new Microlepidoptera. Can. Entomol. 61: 
266-271 (Phlyctaenia sheppardi McDunnough [=Udea sheppardi (McD.))). 

McDUNNOUGH, J. 1938. An apparently new Eupithecia from eastern North America 
(Geometridae: Lepid.). Can. Entomol. 70:171-173 (Eupithecia sheppardata Mc- 
Dunnough). 

McDUNNOUGH, J. 1956. Microlepidoptera notes and a new species. Am. Mus. Novit. 
1789, 17 pp. (Anchylopera sheppardana McDunnough [=Ancylis sheppardana 
(McD.))). 

WALLEY, G. S. 1941. Some new and little known Canadian Ichneumonidae (Hyme- 
noptera). Can. Entomol. 73:164-170 (Allocota sheppardi Walley). 


PAPER ABOUT A. C. SHEPPARD 


VICKERY, V. R. 1969. A. C. Sheppard, the last of a vanishing breed? Bull. Entomol. Soc. 
Canada 1(2):20-21. 


V. R. VICKERY, Emeritus Curator, Lyman Entomological Museum and Research 


Laboratory, Macdonald College, McGill University, Ste-Anne de Bellevue, Québec H9X 
1C0, Canada. 


Date of Issue (Vol. 41, No. 3): 7 October 1987 








EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DouGLas C. FERGUSON, 
ROBERT C. LEDERHOUSE, THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Categories of contributions are Articles, General Notes, Technical Com- 
ments, Book Reviews, and Obituaries. Short manuscripts concerning new state records, 
current events, and notices should be sent to the News, June Preston, Editor, 832 Sunset 
Drive. Lawrence, Kansas 66044 U.S.A. Journal contributors should prepare manuscripts 
according to the following instructions. 

oe An informative rather than descriptive abstract should precede the text of 
Articles. 

Key Words: Up to five key words or terms not in the title should accompany Articles, 
General Notes, and Technical Comments. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely 
double-spaced, employing wide margins, on one side only of white, 8% x 11 inch paper. 
Titles should be explicit and descriptive of content, but must be as short as possible. The 
first mention of a plant or animal in the text should include the full scientific name with 
author. Measurements should be given in metric units; times should be given in terms 
of the 24-hour clock (0930 h, not 9:30 AM). Underline only where italics are intended. 

Literature Cited: References in the text of Articles should be given as Sheppard (1959) 
or (Sheppard 1959, 196ia, 1961b) and listed alphabetically under the heading LITERATURE 
CITED, in the following format without underlining: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In General Notes, references should be given in the text as P. M. Sheppard (1961, Adv. 
Genet. 10:165-216) or (Sheppard, P. M., 1961, Sym. R. Entomol. Soc. London 1:23-30) 
without underlining. 

Illustrations: Only half of symmetrical objects such as adults with wings spread should 
be illustrated, unless whole illustration is crucial. Photographs and drawings should be 
mounted on stiff, white backing, arranged in the desired format, allowing (with particular 
regard to lettering) for reduction to fit a Journal page. Illustrations larger than 8% x 11 
inches are not acceptable and should be reduced photographically to that size or smaller. 
The author’s name, figure numbers as cited in the text, and an indication of the article's 
title should be printed on the back of each illustration. Figures, both line drawings and 
halftones (photographs), should be numbered consecutively in Arabic numerals. The term 
“plate” should not be employed. Figure legends must be typewritten, double-spaced, on 
a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, 
with a separate paragraph devoted to each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical lines as well as vertical writing should be avoided. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer's errors. Excessive author’s changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany the proofs. 

Page charges: For authors supported by institutions, page charges are $20 per Journal 
page. For unsupported authors, page charges are $10 per Journal page with a $50 
maximum. Authors of Book Reviews and Obituaries are exempt from page charges. 
Authors unable to pay page charges for reasons such as difficulties with foreign exchange 
should apply to the editor for free publication. Four free publication slots are available 
annually in the Journal. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A 





CONTENTS 


PATTERNS OF OVIPOSITION IN HEMILEUCA LUCINA (SATURNI- 
IDAE). M. Deane Bowers & Nancy E. Stamp 


PREDATION BY ANOLIS LIZARDS ON BATTUS PHILENOR RAISES 
QUESTIONS ABOUT BUTTERFLY MIMICRY SYSTEMS. Francois 
J. Odzndaal, Mark D. Rausher, Betty Benrey & Juan Nunez- 
Farfan 0. 
NEW RECORDS OF BUTTERFLIES FROM THE WEST INDIES. Albert 
Schwartz, Fernando L. Gonzalez t Rose M. Henderson _. 
A NEW SPECIES OF GRETCHENA (TORTRICIDAE) INJURIOUS TO 
PLANTED NEOTROPICAL WALNUT. William E. Miller 


INJURY AND BIOLOGY OF THE CLEARWING BORER SYNANTHEDON 
KATHYAE ON HOLLY. G. M. Ghidiu, L. Vasvary, T. D. Eich- 
lin ds J. D. Solomon <3 

THE ROLE OF NECTAR SOURCE DISTRIBUTION IN HABITAT USE AND 
OVIPOSITION BY THE TIGER SWALLOWTAIL BUTTERFLY. 
David W. Grossmueller & Robert C. Lederhouse 


GENERAL NOTE 


Penstemon digitalis (Scrophulariaceae), a new food plant record for Haploa 
confusa (Arctiidae). Richard L. Lindroth 


Book REVIEWS 
Mexican Lepidoptera: Eurytelinae I. Sanford R. Leffler ce teeseeeeeeeeeee 
Noctuelles et Géométres d Europe, Volume II. Jeremy Holloway 


OBITUARY 
Arthur Charles Sheppard (1902-1987). V.R. Vickery ccc , 


131 


141 


145 


151 


154 


159 


170 


aS ee fet, 


Volume 41 1987 Number 4 


ISSN 0024-0966 


JOURNAL 


of the 


_ LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 





5 January 1988 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


JERRY A. POWELL, President JEAN-FRANCOIS LANDRY, Vice 
DouGLAS C. FERGUSON, Immediate Past President 

President ATUHIRO SIBATANI, Vice 
JACQUELINE Y. MILLER, Vice President President 
RICHARD A. ARNOLD, Secretary JAMES P. TUTTLE, Treasurer 


Members at large: 


MIRNA M. CASAGRANDE M. DEANE BOWERS JULIAN P. DONAHUE 
EDWARD C. KNUDSON RICHARD L. BROWN JOHN E. RAWLINS 
FREDERICK W. STEHR PAUL A. OPLER Jo BREWER 


The object of the Lepidopterists’ Society, which was formed in May 1947 and for- 
mally constituted in December 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to fa- 
cilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $25.00 
Student members—annual dues $15.00 
Sustaining members—annual dues $35.00 
Life members—single sum $350.00 
Institutional subscriptions—annual $40.00 


Send remittances, payable to The Lepidopterists’ Society, to: James P. Tuttle, Treasurer, 
3838 Fernleigh Ave., Troy, Michigan 48083-5715, U.S.A.; and address changes to: Julian 
P. Donahue, Natural History Museum, 900 Exposition Blvd., Los Angeles, California 
90007-4057 U.S.A. For information about the Society, contact: Richard A. Arnold, Sec- 
retary, 50 Cleaveland Rd., #3, Pleasant Hill, California 94523-3765, U.S.A. 


» 


To obtain: 

Back issues of the Journal and News (write for availability and prices); The Com- 
memorative Volume ($10.00; $6.00 to members, postpaid); A Catalogue/Checklist of 
the Butterflies of America North of Mexico (clothbound $17.00, $10.00 to members; 
paperbound $8.50, $5.00 to members): order (make remittance payable to “The Lepi- 
dopterists’ Society”) from the Publications Coordinator, Ronald Leuschner, 1900 John 
St., Manhattan Beach, California 90266-2608, U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly for 
$40.00 (institutional subscription) and $25.00 (active member rate) by the Lepidopterists’ 
Society, % Los Angeles County Museum of Natural History, 900 Exposition Blyd., Los 
Angeles, California 90007-4057. Second-class postage paid at Los Angeles, California and 
additional mailing offices. POSTMASTER: Send address changes to the Lepidopterists’ 
Society, % Natural History Museum, 900 Exposition Blvd., Los Angeles, California 90007- 
4057. 


Cover illustration: Semilooping larva of the strange noctuid Phyprosopus callitrichoides 
on Smilax. Sketch by Mark Klingler, Carnegie Museum of Natural History. Suggested 
by John E. Rawlins. 


JOURNAL OF 


Tue LeEepipoprTreRIstTs’ SOCIETY 








Volume 41 1987 Number 4 











Journal of the Lepidopterists’ Society 
41(4), 1987, 173-186 


THE BIG SHIFT: NABOKOVI FROM ATALOPEDES 
TO HESPERIA (HESPERIIDAE) 


JOHN M. BURNS 


Department of Entomology, NHB 169, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560 


ABSTRACT. The orange and brown skipper Atalopedes nabokovi, described by Bell 
and Comstock in 1948 and indigenous to xeric lowland thorn scrub of Hispaniola, is 
actually a large and stunning species of Hesperia with no respect for the classically 
Holarctic distribution of that genus. Characters of the male and female genitalia are 
critical both in delimiting the sister genera Atalopedes and Hesperia and in finding the 
sister of nabokovi. (I compared more than 150 KOH-dissections in these two genera.) 
Though highly distinct, Hesperia nabokovi is genitalically (and ecologically) closest to H. 
meskei of the southeastern United States. Genitalic characters, generally so useful in 
differentiating species, are also exceptionally valuable at the generic level in skippers. 
Bell and Comstock, who figured the male genitalia of H. nabokovi, must have been misled 
by the West Indian origin of this skipper and by the large, dark stigma of the male— 
but even that stigma clearly belongs to Hesperia, not Atalopedes. 


Additional key words: genitalia (male and female), stigma, Hesperia meskei, His- 
paniola, variation. 


Our taxonomy can be wrong where we least expect it. In the course 
of reviewing the small genus Atalopedes before adding a couple of 
skippers to it (Burns in prep.), I finally obtained specimens of the species 
endemic to Hispaniola. From the figure of male genitalia in the original 
description (Bell & Comstock 1948), this species had already struck me 
as quite the most primitive member of Atalopedes—the first to arise 
after the sister genera Hesperia and Atalopedes split. Except for the 
fact that the dorsal edge of the valva was almost uniform in height 
throughout its length, instead of humped near the middle and lower 
at either end, the genitalia looked like those of Hesperia; but certain 
critical features of the uncus and the penis did not show in the lateral 
view provided. And then there was the rest of the animal to wonder 
about. The moment I saw it, it bothered me: though the facies could 
fit Atalopedes or Hesperia, the stigma belonged to Hesperia. Still, “the 


174 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


genitalia are the best place to start” (Burns 1985:3). Sure enough, after 
perusing the first dissection of each sex and the comparative figures in 
MacNeill (1964), I knew that nabokovi is a species of Hesperia. 

This hurts the common generalization “Hesperia is Holarctic” (Klots 
(1951, Evans 1955, MacNeill 1964, 1975). Now we must say that Hes- 
peria is Holarctic and Hispaniolan, which is less tidy but more allit- 
erative—and a healthy reminder that, despite present distributions, 
Hesperia does not have to be northern in ultimate origin. Of course, 
its tropical occurrence in the heart of the West Indies need not connote 
some enormous ecologic leap. The southernmost eastern species of Hes- 
peria on the continent, H. meskei (Edwards), whose range includes not 
only peninsular Florida but also Florida Keys, inhabits such hot, dry 
communities as pine woods or barrens and oak scrub or woodland or 
savanna (McGuire 1982, Burns unpubl.). The Hesperia on Hispaniola 
occupies most of the xeric lowland thorn scrub (A. Schwartz pers. 
comm.). 


Hesperia nabokovi (Bell & Comstock), new combination 


Atalopedes nabokovi Bell & Comstock (1948:19). Evans (1955:339); Riley (1975:186). 


The long, verbal original description (of one male and one female 
from Haiti) dealt with little besides facies and its considerable sexual 
dimorphism. Description of genitalia was confined to the male and, at 
that, to a figure (left lateral view), except for a single (under)statement: 
“The male genitalia show specific differences from those of [Atalopedes] 
campestris [(Boisduval)], the unci and the terminations of the claspers 
being different in the two species.’’ Actually, the genitalia show generic 
differences—in both sexes—from those of Atalopedes and thoroughly 
fit the Hesperia mold. 

Rather than conventionally redescribe H. nabokovi, I will discuss 
selected characters in connection with its proper generic placement, its 
situation within Hesperia, its peculiarities, and its variability. Genitalic 
terminology largely follows MacNeill (1964). 


Male Genitalia (Figs. 1-7) 


In H. nabokovi, as in half the Nearctic species of Hesperia (MacNeill 
1964), the uncus forms a slender, caudally produced, medial beak— 
the fine, median dorsoventral cleft at its apex becoming relatively long 
(Fig. 1). To correspond with the uncus, the paired underlying gnathos 
lengthens (Figs. 1, 2). As in Hesperia generally, the valva ends in two, 
more or less prominent, pointed dorsal teeth whose bases are connected 
on the outer surface of the valva by a smooth and conspicuous U-shaped 
edge (Fig. 2). From the distal tooth, an irregular dentate edge—the 


VOLUME 41, NUMBER 4 75 





Fics. 1-8. Male genitalia of Hesperia nabokovi from 4 km E El Limon, ca. 185 m 
(600 ft), Independencia, Dominican Republic, 16 October 1988, A. Schwartz (genitalic 
dissection no. X-2196). 1, Tegumen, uncus, and gnathos in dorsal view; 2, Complete 
genitalia (minus right valva) in left lateral view; 3, Penis and juxta in dorsal view. These 
figures show certain structures in parallel alignment at two different angles, 90° apart, so 
as to convey form in three dimensions. 


“inner serration”’ of MacNeill—runs ventrad and cephalad, medial to 
the proximal tooth (Fig. 2). Again in the Hesperia pattern, the penis 
bears a small, bidentate cornutus distally in the dorsal vesica and a 
larger, bidentate projection left-laterally at its distal end (Figs. 2, 3). 
This projection MacNeill (1964) called the rostellum; but I prefer the 
more suggestive loose synonym titillator (Tuxen 1956), especially on 
account of its striking hypertrophy in H. nabokovi (Fig. 3). The penis 
is no longer than the rest of the intact genitalia from the anterior tip 
of the saccus to the posterior tips of the uncus and valvae (Fig. 2). As 
in other Hesperia (figures in Skinner & Williams 1924b and Lindsey 
et al. 1931), paired prongs projecting anteriorly from the anterior end 
of the juxta are long and delicate (Figs. 2, 3). 


176 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


In Atalopedes, by contrast, the uncus is stubby; the gnathos, moderate 
to vestigial; and the valva, elongate, with neither the inner serration of 
all species of Hesperia nor the decided dorsal hump of virtually all of 
them (so that, in lateral view, top and bottom of the valva are about 
parallel). The penis—which is much longer than the rest of the geni- 
talia—has either two, relatively elaborate, multidentate cornuti or none 
at all. The paired prongs projecting anteriorly from the anterior end 
of the juxta are comparatively short and stout (Burns in prep.). 

With respect to these many and various characters, H. nabokovi 
resembles Atalopedes only in having an elongate valva without a dorsal 
hump. 

Owing to its long, beaked uncus and long gnathos, H. nabokovi goes 
with an array of Hesperia species treated by MacNeill (1964) as “the 
Metea species group,” on the one hand, and as “species of uncertain 
affinities,’ on the other. The former includes H. attalus (Edwards), H. 
metea Scudder, and H. viridis (Edwards); the latter, H. meskei, H. 
dacotae (Skinner), H. lindseyi (Holland), H. sassacus Harris, H. miri- 
amae MacNeill, and H. nevada (Scudder). These latter species “‘con- 
stitute a very diverse and possibly unnatural assemblage. The morpho- 
logical divergence apparent between these species is of a magnitude 
found between species groups elsewhere in the genus” (MacNeill 1964: 
57). 

The last three species (sassacus, miriamae, and nevada)—which are 
all adapted to cold—differ from nabokovi on various genitalic counts. 
What may be the most critical involves the point where the paired 
gnathos joins the tegumen—the “gnathos insertion” of MacNeill—which 
is much farther forward in these than in any other species of Hesperia, 
including nabokovi. Moreover, the uncus beak is exceptionally long in 
both sassacus and nevada; and the cleft in its apex is lengthened in 
sassacus and eliminated in nevada (a state unique in the genus). 

The next two species (dacotae and lindseyi) also differ from nabokovi 
in many respects, not least of which is a tendency of the titillator to 
enlarge more posteriad than laterad and to develop additional teeth. 
This is carried to an unparalleled extreme in dacotae, whose hypertro- 
phied titillator extends straight back beyond the distal penile opening 
to yield the longest penis in the genus. 

On the basis of shape and length of tegumen and beaked uncus, 
length of the uncal cleft, and length of gnathos and level of its insertion, 
nabokovi most nearly resembles meskei and the species of the Metea 
group (attalus, metea, and viridis). These last three species, though 
grouped, “are not very closely related’’ to one another (MacNeill 1964: 
151); and their nearest (but distant!) ally may be meskei: “this species 


VOLUME 41, NUMBER 4 VE 


is not placed easily within any of the preceding groups of species, 
although it perhaps resembles the Metea group more closely than it 
resembles any of the [other] species [of uncertain affinities!’ (MacNeill 
1964:157). Hesperia nabokovi apparently belongs in a section of the 
genus comprising scattered remnant species. MacNeill (1964:13, pers. 
comm.) considers them some of the oldest in Hesperia. To generalize 
from present geographic distributions, these species are especially tol- 
erant of heat. 

Having roughly rooted H. nabokovi, I must hasten to emphasize that 
it not only lacks close relatives but flaunts more than its share of genitalic 
idiosyncrasies. The vinculum is uniquely narrow where it joins the 
tegumen—a feature best seen in lateral view (Fig. 2). The tegumen is 
uniquely long in the zone of dense bristles, which extends back to about 
the uncus and the gnathos insertion (Figs. 1, 2). The valva is elongate, 
with little or no hump on its dorsal edge (Fig. 2). (The nearest approach 
in other Hesperia to this anomalous humpless condition is in dacotae.) 
In dorsal view, at or a little beyond the level of the distal end of the 
juxta, the valvae are not characteristically “plump” as they are in other 
Hesperia. The broadly rounded distal end of the valva protrudes ap- 
preciably caudad of the distal tooth—more so than in any other Hes- 
peria (Fig. 2). Last but not least, the hypertrophied, almost rectangular, 
heavily sclerotized titillator expands to the left from the distal end of 
the penis like a small, stiff flag on a stout pole (Fig. 3), the two titillator 
teeth typical of other Hesperia becoming the outer corners of the flag. 

In their revisions of Hesperia, both Lindsey (1942) and MacNeill 
(1964) stressed the extraordinarily high levels of individual variation 
encountered. Naturally enough, such variation can loosen the genitalia 
as well as the external phenotype. Among just three males of H. nabokovi 
at hand, the overall shape of the tegumen plus uncus varies noticeably. 
From what seems the most nearly average condition (Fig. 1), one male 
departs in the direction of a malformed tegumen whose transition to 
uncus is abruptly concave (Fig. 4); the other, in a svelte direction— 
decidedly longer and narrower (Fig. 5). (Much of the apparent variation 
in the gnathos, however, simply stems from the mobility of its two 
separate arms.) Again, in these three males, the proximal tooth of the 
valva bends inward as little as 20° to as much as 90°, while the distal 
tooth stays upright (Figs. 2, 6, 7). Because a much-bent tooth vanishes 
in lateral view (Fig. 7), a casual observer might fail to see the config- 
uration so characteristic of Hesperia. Inbending even involves some of 
the dorsal edge of the valva anterior to the proximal tooth; and, curiously 
enough, a slight asymmetry crops up, with the left tooth (and rim) 
bending more than the right. As usual in Hesperia, the inner serration 


178 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


6 








Fics. 4-7. Male genitalia of Hesperia nabokovi from 11.5 and 12 km ESE Canoa, 
Barahona, Dominican Republic: 4, 6, 31 July 1982, F. Gali (X-2195); 5, 7, 7 August 1986, 
A. Schwartz (X-2197). 4, 5, Tegumen, uncus, and gnathos in dorsal view, with the gnathos 
insertion indicated; 6, 7, Left valva in lateral view. 


varies in detail (Figs. 2, 6, 7). One male of nabokovi starts to express 
the typical Hesperia hump on the dorsal margin of the valva (Fig. 7). 
The rectangular titillator expands a bit ventrad as well as laterad in 
two out of three males. 

Even though the angle of the proximal tooth varies greatly in na- 
bokovi, its orientation relative to the distal tooth (within a single valva) 
has taxonomic merit. In the nine males of H. meskei examined, the 
proximal tooth bends inward about 15 to 30° while the distal tooth 
stands erect. Essentially, then, in both nabokovi and meskei the proximal 
tooth is medially inclined whereas the distal tooth is about vertical so 
that, in posterior view, their paths seem to meet or cross. On the con- 
trary, in H. metea, H. viridis, and H. attalus (the Metea group), both 
the proximal tooth and the distal tooth are medially inclined—and to 
similar degrees—so that, in posterior view, they look about parallel. 
Moreover, in lateral view, despite ample variation, the proximal and 
distal teeth are relatively far apart in nabokovi (Figs. 2, 6, 7) and meskei 
but close together in the species of the Metea group (especially metea 
and attalus); the proximal tooth is shorter than, or, at most, equal to, 
the distal tooth in nabokovi (Figs. 2, 6, 7) and meskei but taller than 
the distal tooth in the Metea group; and the inner serration, at its 
proximal end, is not toothed in nabokovi (Figs. 2, 6, 7) and modestly 
toothed in meskei but strongly toothed in the Metea group. Again, in 
lateral view, the U on the outer surface of the valva connecting the 
bases of the proximal and distal teeth is so deep in nabokovi (Figs. 2, 
6, 7) and meskei that it exposes most of the inner serration but so shallow 


VOLUME 41, NUMBER 4 179 


in the Metea group that it hides it. All things considered, nabokovi is 
closest to meskei. 


Female Genitalia (Figs. 8, 9) 


Both Hesperia and Atalopedes reflect a broader pattern in which the 
sterigma and the ductus bursae are sclerotized while the corpus bursae 
is membranous. 

In H. nabokovi, as in Hesperia generally, the outline of the lamella 
postvaginalis is roughly rectangular in ventral view (Fig. 8), the ductus 
bursae is angled to the left (Fig. 8), a singular expansion of the ductus 
bursae—corresponding to the “caudal chamber” of MacNeill (1964)— 
is asymmetrically developed on the right (Figs. 8, 9), the ductus bursae 
is still sclerotized not only where the ductus seminalis joins but also well 
cephalad of that level (Fig. 9), and the corpus bursae is roughly spherical 
(Figs. 8, 9). 

In Atalopedes a sclerotized midventral prong (short to long, according 
to species) projects caudad, or caudad and ventrad, from the posterior 
part of the lamella postvaginalis. The rest of the lamella postvaginalis 
comprises (1) midventral sclerotization that carries the dorsal wall of 
the ductus bursae back to the base of the prong, and (2) a closely flanking 
pair of large, smooth “plates” (variously ovate to comma- or kidney- 
shaped) that spread dorsad and laterad. Depending on the species, 
sclerotization of these plates may be strong throughout or so weak that 
only their medial margins show. In any case, seen ventrally, the lamella 
postvaginalis as a whole does not suggest a rectangle. The ductus bursae— 
which is neither angled nor asymmetrically developed—does not enter 
the corpus bursae directly but by way of a dorsal jog. The ductus 
seminalis joins the ductus bursae at this jog, which is membranous 
(coming right after the sclerotized portion of the ductus bursae). The 
elongate corpus bursae looks like a sausage (Burns in prep.). 

The female genitalia buttress those of the male in suggesting that H. 
meskei is the nearest living relative of H. nabokovi. In both species the 
sterigma is unusually simple and lightly sclerotized while the caudal 
chamber of the ductus bursae is unusually expanded and heavily scler- 
otized. More explicitly, the lamella postvaginalis looks squarish and 
relatively flat—without bold features of relief—in ventral view. Its light 
sclerotization is mainly peripheral (distal and/or lateral), leaving an un- 
to barely sclerotized central zone. The lamella antevaginalis is fully 
membranous (it is partly sclerotized in other species of Hesperia— 
almost always conspicuously and always perceptibly). The caudal cham- 
ber is exceptionally large and flat, expanding far laterad (to the right) 
but not, or not far, ventrad. 

Despite some broad similarity of the caudal chamber in H. nabokovi 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 8,9. Female genitalia of Hesperia nabokovi from 1 km SE Monte Cristi, 0 m, 
Monte Cristi, Dominican Republic, 15 May 1986, F. L. Gonzalez (X-2194). 8, Sterigma 
and bursa copulatrix in ventral view; 9, The same, plus part of the ductus seminalis, in 
right lateral view. 


and H. meskei, the ductus bursae as a whole differs a lot. In nabokovi 
it is short; even apart from the caudal chamber, it is wide; and ventrally 
it is fully sclerotized as far caudad as the start of the lamella postvagi- 
nalis. Conversely, in meskei it is long; apart from the caudal chamber, 
it is narrow; and ventrally it is not sclerotized to the level of the lamella 
postvaginalis. The most striking difference, however, involves such total 
incorporation of the caudal chamber into the ductus bursae in nabokovi 
that the chamber is no longer the caudalmost element of the ductus 
(Figs. 8, 9)—a condition unique in the entire genus (compare figures 
in MacNeill 1964:194, 218-221). 

Although the morphologically simpler female genitalia are less glar- 
ingly variable than those of the male, they are still highly individual 
in the three females examined, especially in the outline of the heavily 
sclerotized caudal chamber, the length and angle of the ductus bursae, 
and the outline of the lamella postvaginalis. To illustrate, the caudal 
chamber is somewhat rounded in the female drawn (Fig. 8) but more 


VOLUME 41, NUMBER 4 181 





Fic. 10. Stigma on the dorsal left primary of the Hesperia nabokovi male whose 
genitalia appear in Figs. 4 and 6. 


rectangular in the other two females, in one of which it is also longer 
and narrower (as is the entire ductus bursae). Again, in these three 
females, the midventral third of the posterior margin of the lamella 
postvaginalis is conspicuously concave, slightly so (Fig. 8), or slightly 
convex. 


Stigma (Fig. 10) 


When Bell and Comstock (1948:20) described nabokovi in Ata- 
lopedes, they said “‘the stigma is relatively very large but of the form 
characteristic of the species in this genus’; and when Riley (1975:186) 
treated nabokovi in his guide to West Indian butterflies, he said “‘sex 
brand as in A. mesogramma [(Latreille)].’’ Not so: the stigma of nabokovi 
carries the Hesperia stamp (which has been well characterized by 
MacNeill 1964:49, 57, 194). 

In H. nabokovi, the usual parts are present and in place (Fig. 10). 
Most telling are the two rows of large, wide, silvery-gray scales enclosing 
the dustlike microandroconial mass to form a conspicuous, gentle arc. 
Flanking this centerpiece costally and basally are the narrow, dark 
apical and lower brush patches; and flanking it outwardly is the broad, 
dark poststigmal patch. 

The microandroconial mass is dark gray, as it is in most species of 
Hesperia, not yellow, as it is in the species of the Leonardus group. 


182 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Facies (Figs. 11-15) 


Essentially, these skippers are orange and brown. 

Above, males are mostly bright orange, with a dark outer margin on 
the primary, a narrow (linear) dark outer margin plus a broad dark 
costal margin on the secondary, and very narrowly darkened veins in 
both wings (Fig. 11). The large, dark, central stigma dominates the 
broadly orange primary—and, for that matter, the entire dorsal aspect 
(Fig. 11). Two small, orange subterminal spots vaguely mark the inner 
edge of the dark margin in spaces 4 and 5 of the primary. Below, an 
unworn male looks mostly dull orange, except for the narrowest of 
linear dark costal and outer margins on both wings and a very dark 
basal area (easily hidden) on the primary (Fig. 13). Much of the “dull- 
ness” stems from a scaling of orange upon brown: loss of overscaling 
in worn males reveals a brown ground color across the apex of the 
primary and over all of the secondary except space 1b and an adjacent 
strip of space lc (Fig. 12). The orange overscaling largely obscures pale 
orange subterminal spots in spaces 4 and 5 and apical spots in spaces 
6 and 7 (sometimes also 8) of the primary as well as basal spots and 
spots of the macular band of the secondary (compare worn and unworn 
undersides in Figs. 12 and 18, respectively). (Spot terminology follows 
Lindsey 1942 and especially MacNeill 1964:49, 194.) 

As in Hesperia and related genera generally, brown coloring develops 
at the expense of orange in females so as to yield darker skippers. Both 
above and below, females of H. nabokovi show more pattern, more 
spots than do males (Figs. 14, 15). Spots are opaque. Above, they are 
orange. But below, the apical and subterminal spots of the primary and 
the spots of the secondary are white. (Secondary spots include basal 
spots in the cell and space 7 plus the spots of the macular band, which 
may extend from space lc to space 7 when maximally expressed.) 
Moreover, in unworn females, dark scales ring the white spots of the 
macular band (Fig. 14), while the overscaling (which covers the same 
brown ground as in males) has a greenish cast—all to stunning effect. 

Although Hesperia is a notoriously difficult genus, H. nabokovi should 
not be confused with any other species. 

The original description (Bell & Comstock 1948) accurately char- 
acterized in many words the facies of a single male and female. The 
only figures of facies to date are black-and-white drawings (Riley 1975: 
186) of those very same specimens. When I lent the holotype male from 
the Museum of Comparative Zoology, Harvard University, to Riley in 
1974 for illustration, its true generic identity escaped me; but, for what 
it’s worth, I can honestly say that I have seen and held the holotype of 
nabokovi. 


VOLUME 41, NUMBER 4 183 





de 14 15 


Fics. 11-15. Facies of Hesperia nabokovi (all x1): 11, 15, dorsal views; 12-14, ventral 
views. 11, 12, The worn male whose genitalia appear in Figs. 4 and 6 and stigma, in 
Fig. 10; 13, The unworn male whose genitalia appear in Figs. 5 and 7; 14, 15, An 
unworn female from 4 km SE Monte Cristi, Monte Cristi, Dominican Republic, 18 October 
1983, J. W. Raburn (X-2198). 


Antenna 


In all three males and two of the three females at hand, the nudum 
of the antenna is 8/5; that is, there are 8 bare segments on the main 
mass of the club plus 5 on the apiculus for a total of 13. In the third 
female the nudum is 9/5. 

Evans (1955:300, 301, 317, 338) gave the nudum of Hesperia as 8/4 
and that of Atalopedes as 7/7. This character is more variable and 
more difficult to score than Evans would have you believe. 


Size 
The length (mm) of one primary in the males is 17.1, 17.5, and 18.6; 
in the females, 19.0, 20.0, and 20.1. Bell and Comstock (1948:21) gave 
a primary length of 20 mm for the holotype male and 18 mm for the 
allotype female. 
This is a large species of Hesperia. 


Spatial and Temporal Distribution 


As noted at the outset, H. nabokovi occurs in the xeric lowland thorn 
scrub on Hispaniola, which is extensive. Exact data on the specimens 
available to me, which come from the northwestern and southwestern 
Dominican Republic, appear in figure legends. (The one female not 
specifically cited, genitalic dissection X-2192, has the same data as the 
male in Figs. 1-8.) The holotype and allotype are from Thomazeau 
and Fond Parisien, respectively, both in southeastern Haiti (Bell & 


184 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Comstock 1948). Altogether, these eight specimens represent six dif- 
ferent months—February, May, July, August, September, October— 
indicating that H. nabokovi is multivoltine. 


DISCUSSION 


Why did Bell and Comstock (1948) put nabokovi in Atalopedes and 
not Hesperia, especially with so many relevant genitalic illustrations 
about. Within the preceding quarter century, Skinner and Williams 
(1924a, 1924b) had figured the male genitalia of A. campestris and the 
American species of Hesperia; Lindsey et al. (1931) had reprinted all 
those figures; Lindsey (1942) had newly refigured the male genitalia 
of the entire genus Hesperia; and Comstock (1944) had figured the 
male genitalia of A. mesogramma. Clearly, Bell and Comstock gave 
too little weight to genitalic morphology and far too much to the dark 
color and large size of the poststigmal patch. And (subconsciously, at 
least) they must have thought Haiti too tropical, too insular—altogether 
too outlandish—for a Holarctic genus like Hesperia. Atalopedes, on the 
other hand, had long been known from the West Indies and from 
Central and northern South America in the form of A. mesogramma 
and A. campestris, respectively. 

There was still a lingering reluctance among American skippermen 
to give the genitalia their taxonomic due. To appreciate this, one need 
only study the genitalic figures in Lindsey et al. (1931) for such genera 
as Polites (treated as Talides) on pages 97 and 101 or Atrytone on 
pages 112, 115, and 119: the genitalia of P. verna (Edwards) are in no 
way a variation on the repetitious genitalic theme of other species of 
Polites nor are those of A. arogos (Boisduval & Le Conte) and A. logan 
(Edwards) variations on the different but equally repetitious genitalic 
theme of other species of Atrytone. Eventually, verna was moved to 
the new genus Pompeius and all species of Atrytone except arogos and 
logan, to Euphyes by the Englishman Evans (1955). 

Genitalia deserve all the respect and attention they can get, which 
means, in general, that they should be weighted heavily—and (no mere 
converse) that they should not be used lightly: nowhere are analysis of 
variation and interpretation in context more important. Genitalia may 
be remarkably conservative among species in some genera or complexes 
(even to the point of yielding no diagnostic characters) and yet won- 
derfully differentiated among species in others. What may amount to 
a subtle but real interspecific difference in one instance may be nothing 
more than individual variation in another. And so forth. Every use of 
genitalia in systematics calls for thorough background investigation. I 


VOLUME 41, NUMBER 4 185 


have repeatedly answered questions of individual, geographic, and in- 
terspecific variation in skipper genitalia by dissecting and comparing 
large samples (usually both sexes) from many areas, especially in Eryn- 
nis, Celotes, Atrytonopsis, Autochton, Wallengrenia, and Pyrgus (Burns 
1964, 1970, 1974, 1983, 1984, 1985, unpubl.). 

My small genitalic sample of H. nabokovi (three males, three females) 
looks better in light of the limited and isolated geographic range of this 
species. And the considerable individual variation evident in this sample 
looks minor next to the grand and pervasive genitalic divergence that 
exists between nabokovi and all other species of Hesperia. 

I have stressed from the start that, in both sexes, the genitalia of 
nabokovi—despite their distinctive attributes—are assuredly those of 
Hesperia. In this connection I note that, within the set of diverse species 
having a long, beaked uncus and so including nabokovi, species ap- 
parently not closest to it sometimes express character states reminiscent 
of it. For example, H. dacotae tends to approximate the elongate, 
humpless valva; and H. nevada, the hypertrophied titillator, as well as 
the simple, lightly and peripherally sclerotized lamella postvaginalis 
coupled with the caudal chamber expanding substantially to the right. 
Such similar genitalic tendencies will most likely surface independently 
among species that are still genetically similar. 

However that may be, the magical writer and lepidopterist Vladimir 
Nabokov would doubtless have enjoyed this switch to the type-genus 
of the family Hesperiidae. 


ACKNOWLEDGMENTS 


Thanks to J. Y. Miller for steering me to Albert Schwartz in my quest for H. nabokovi; 
to Schwartz for generously lending three pairs of this skipper for genitalic dissection; to 
B. A. B. Venables for KOH-dissecting these and 102 other specimens of Hesperia, as well 
as 32 specimens of Atalopedes; to Young Sohn for elegantly drawing genitalia and 
mounting photographs; to V. E. Krantz for photographing the half animal and its stigma; 
and, above all, to C. D. MacNeill for making such sense of the stubborn genus Hesperia 
in the first place. 

MacNeill and S. S. Nicolay kindly reviewed the manuscript. 


POSTSCRIPT 


Thanks to Kurt Johnson and F. H. Rindge, I saw three more females of Hesperia 
nabokovi, and their dissected genitalia, in October 1987: the allotype (from the American 
Museum of Natural History, New York), which was taken in 1922 at an elevation of 
about 60 ft (18 m) in Haiti, plus two females taken recently at even lower elevations in 
the desert around Cabo Rojo, Pedernales, Dominican Republic. Variation in the genitalia 
is conspicuous, matching what I have already described. With respect to facies, both Cabo 
Rojo females have “unusually” small spots below; and above, one female is remarkably 
orange and hence bright, while the other is as dark as the now faded allotype once was. 
Predictably, the larger sample pushes variation toward the rampant state so common in 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hesperia. Nudum counts are again 8/5 and 9/5; primary lengths, another 20.0 mm and 
a whopping 21.5 mm. 


LITERATURE CITED 


BELL, E. L. & W. P. Comstock. 1948. A new genus and some new species and subspecies 
of American Hesperiidae (Lepidoptera, Rhopalocera). Am. Mus. Novitates No. 1379. 
23 pp. 

BuRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. 
Publ. Entomol. 37:1-217. 

1970. Secondary symmetry of asymmetric genitalia in males of Erynnis funeralis 

and E. propertius (Lepidoptera: Hesperiidae). Psyche 77:480-435. 

1974. The polytypic genus Celotes (Lepidoptera: Hesperiidae: Pyrginae) from 

the southwestern United States and northern Mexico. Psyche 81:51-69. 

1983. Superspecies Atrytonopsis ovinia (A. ovinia plus A. edwardsi) and the 

nonadaptive nature of interspecific genitalic differences (Lepidoptera: Hesperiidae). 

Proc. Entomol. Soc. Wash. 85:335-358. 

1984. Evolutionary differentiation: Differentiating gold-banded skippers—Au- 

tochton cellus and more (Lepidoptera: Hesperiidae: Pyrginae). Smithsonian Contrib. 

Zool. No. 405. 38 pp. 

1985. Wallengrenia otho and W. egeremet in eastern North America (Lepi- 
doptera: Hesperiidae: Hesperiinae). Smithsonian Contrib. Zool. No. 423. 39 pp. 
Comstock, W. P. 1944. Insects of Porto Rico and the Virgin Islands—Rhopalocera or 
butterflies. New York Acad. Sci., Scientific Survey of Porto Rico and the Virgin Islands 
12:421-622. 

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the classifi- 
cation and nomenclature adopted in the British Museum (Natural History). Part IV. 
Hesperiinae and Megathyminae. British Museum, London. 499 pp., pls. 54-88. 

K.Lots, A. B. 1951. A field guide to the butterflies of North America, east of the Great 
Plains. Houghton Mifflin Co., Boston. 349 pp., 40 pls. 

LinpsEy, A. W. 1942. A preliminary revision of Hesperia. Denison Univ. Bull. J. Sci. 
Lab. 37:1-50, pls. 1-6. 

LINDsEY, A. W., E. L. BELL & R. C. WILLIAMS JR. 1931. The Hesperioidea of North 
America. Denison Univ. Bull., J. Sci. Lab. 26:1-142. 

MACNEILL, C. D. 1964. The skippers of the genus Hesperia in western North America 
with special reference to California (Lepidoptera: Hesperiidae). Univ. Calif. Publ. 
Entomol. 35:1-230. 

1975. Family Hesperiidae, pp. 423-578. In Howe, W. H. (ed.), The butterflies 
of North America. Doubleday & Co., Inc., Garden City, New York. 

McGuire, W. W. 1982. Notes on the genus Hesperia in Texas: Temporal and spatial 
relationships. Bull. Allyn Mus. No. 78. 21 pp. 

RILEY, N. D. 1975. A field guide to the butterflies of the West Indies. Collins, London. 
224 pp., 24 pls. 

SKINNER, H. & R. C. WILLIAMS JR. 1924a. On the male genitalia of the Hesperiidae of 
North America, Paper V. Trans. Am. Entomol. Soc. 50:141-156. 

1924b. On the male genitalia of the Hesperiidae of North America, Paper VI. 
Trans. Am. Entomol. Soc. 50:177—208. 

TUXEN, S. L. (ed.) 1956. Taxonomist’s glossary of genitalia in insects. Ejnar Munksgaard, 
Copenhagen. 284 pp. 























Received for publication 27 June 1987; accepted 11 August 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 187-194 


LITODONTA HYDROMELI HARVEY (NOTODONTIDAE): 
DESCRIPTION OF LIFE STAGES 


S. J. WELLER 
Department of Zoology, University of Texas, Austin, Texas 78712 


ABSTRACT. Litodonta hydromeli defense structures, mandibular morphology, feed- 
ing behaviors and setal morphology differ among instars. The third abdominal SV1 seta 
is twice as long in the first than in later instars. Later instars have numerous, scattered 
secondary setae. First instars skeletonize leaves with forklike mandibles. Subsequent instars 
ingest entire leaves, and their mandibles have a thin, ridged, cutting edge. First instars 
possess elaborate prothoracic scoli and dorsal spines which are progressively reduced in 
later instars as a prothoracic gland develops. Larvae spray an acidic substance which 
deters ants. Litodonta hydromeli feeds on Bumelia species (Sapotaceae) and developed 
indoors from egg to prepupa in six weeks. Adults reproduced readily in captivity. 


Additional key words: Heterocampini, ontogeny, immatures, defense gland. 


Notodontid adults have been described as medium-sized, noctuidlike 
moths (Forbes 1948), and probably have been neglected in ecological 
and systematic studies due to their plain appearance. The caterpillars, 
however, are notable for their bizarre shapes and developmental changes 
(ontogeny) (Packard 1895, Holloway 1983). As early as 1895, Packard 
recommended studies of larval ontogeny for systematic purposes. Al- 
though coloration and gross morphology have been described for several 
notodontid immatures (Peterson 1948, Dyar 1904, Godfrey 1984, God- 
frey & Appleby 1987), detailed information of setal arrangement and 
larval ontogeny is lacking for most species. 

Litodonta hydromeli Harvey is a north temperate representative of 
the tribe Heterocampini, and the species occurs in Florida, Texas, Okla- 
homa and Missouri (Harvey 1876b, Kimball 1965, Stephen Passoa pers. 
comm.). Harvey (1876a, 1876b) and Packard (1895) gave general de- 
scriptions of adult habitus. Litodonta hydromeli is figured in Holland 
(1908, plate 39, fig. 20), Kimball (1965, plate 20, fig. 3), and Packard 
(1895, plate 5, fig. 16). Dyar (1904) described the eggs and last instar, 
and gave Bumelia angustifolia (Nutt.) (Sapotaceae) as the larval host. 
Like some other Heterocampini, Litodonta hydromeli transforms from 
a spiny first instar to a cryptic fifth instar with a cervical defense gland 
(Herrick & Detwiler 1919, Eisner et al. 1972, Weatherston et al. 1979). 

The purpose of this paper is to detail larval, pupal, and adult mor- 
phology of L. hydromeli, and to discuss changes in larval defense struc- 
tures and feeding biology during development. 


METHODS 


Adults were taken at blacklights at Austin, Texas, in March 1983, 
March 1984, September 1985, and April 1986. Females (culture Nos. 


188 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


W84-102, WE85-25, and WE85-37) were placed in jars and eggs ob- 
tained. I reared cultures at 24°C on the leaves of Bumelia lanuginosa 
(Michx.) and an unidentified Bumelia species in the laboratory. Eggs 
and subsequent immature stages were preserved in 80% ethanol. Third, 
fourth, and fifth instars were killed in simmering water before pres- 
ervation. Shed head capsules were preserved, and some treated with 
10% KOH, dehydrated, and slide mounted for study of chaetotaxy. 
First and second instars were cleared with cold KOH or 2% trisodium 
orthophosphate, then stained with chlorozol black (Kodak or ICN) dis- 
solved in 20% ethanol. Means are followed by the range in parentheses, 
except where ranges do not exist. Setal nomenclature follows Hinton 
(1946). Male and female genitalia, and male appendages were softened 
in KOH, dissected in 40% ethanol, and stained. Preparations were 
mounted in Canada balsam. Voucher specimens are in the National 
Museum of Natural History, Washington, D.C. 


RESULTS 


Litodonta hydromeli is multivoltine at Austin, and appears to be 
multivoltine in Florida also (Kimball 1965). As reported by Harvey 
(1876b), I found at least three broods to occur in March, April, and 
September. The moths overwintered as prepupae in the laboratory. 
Females laid eggs on the leaves and bark of Bumelia. Development 
lasted approximately six weeks from egg to prepupa. Eggs took six to 
seven days to develop. For all stages, intermolt time was approximately 
five days. Prepupae turned reddish, entered a wandering stage that 
lasted one day, burrowed into soil, and formed slight, silk cocoons within 
earthen pupal cells. In nonwintering individuals, time from prepupa 
to eclosion was about 17 days. Even in these, the prepupa did not 
immediately form a pupa within the cell. Emerging adults mated readi- 
ly in captivity, and a second spring brood was obtained. The fall flight 
occurred in late August and early September. 


Description of Stages 


Egg (n = 4). Dome shaped; light blue-green, becoming yellow, then brown, before 
hatching. Diam. 0.99 mm. Chorion with fine, reticulate sculpturing. 

First instar (n = 7). Length 3.1 (2.5-3.7) mm. Head dark brown. Body ground color 
dark reddish brown; with dark brown scoli and spines. Prolegs and ventral areas yellow 
green with red flecks, A10 proleg bright yellow. Head diam. 0.6 mm; height of frons 0.2 
mm; length of epicranial suture 0.2 mm. Chaetotaxy as in Fig. 1. Mandibles with 3 
fingerlike lobes (Fig. 2). Prothorax with dorsal, three-pronged scoli (Fig. 4); scoli rugose, 
sclerotized base extending to anterior margin of mesothorax; XD1 located on midpoint 
of second prong; XD2 on anterior margin of sclerotized base beneath and anterior to 
XD1; D1 between prongs on sclerotized base and D2 ventrad and on margin. Other 
thoracic primary setae as in Fig. 8. D1 of abdominal segments Al—A6, A8 and A10 on 
raised, serrate pinacula; SV1 of A3 extremely long, twice length of other setae, ventrally 


VOLUME 41, NUMBER 4 189 





























Fics. 1-9. Litodonta hydromeli larval morphology. 1, Head chaetotaxy of first instar; 
2, First instar mandible; 3, Fifth instar mandible; 4, Prothoracic scoli of first instar; 5, 
Prothoracic scoli of third instar; 6, Prothoracic scoli of fourth instar; 7, Prothoracic scoli 
of fifth instar; 8, Chaetotaxy of thorax and Al to A4 of first instar; 9, Chaetotaxy of 
thorax and Al to A4 of fifth instar. Scales represent 1 mm unless otherwise indicated. 


directed. Prolegs on A3—A6 well developed, those on A10 reduced. Crochets uniordinal, 
uniserial. Chaetotaxy as in Fig. 8. 

Second instar (n = 4). Length 3.6 (3.1—4.3) mm. Numerous scattered secondary setae 
present in this and all subsequent instars. Coloration similar to first instar. Head diam. 
0.8 (0.8-0.9) mm; height of frons 0.2 (0.2-0.3) mm; length of epicranial suture 0.4 (0.3- 
0.4) mm. Mandibles with ridgelike cutting edge. Prothoracic scoli as in first instar but 
slightly reduced and more stout. D1 of segments Al—A6 and A8 on simple, raised pinacula; 
D1 of Al0 on slightly raised pinacula; SV1 of A3 normal. 

Third instar (n = 5). Length 11.1 mm (n = 2). Head dark brown; body green with 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


sublateral white stripe. Head diam. 1.4 (1.2-1.5) mm; height of frons 0.4 (0.3-0.4) mm; 
length of epicranial suture 0.7 (0.6—-0.8) mm; vertex slightly narrower than in previous 
instars. Prothoracic scoli reduced, but with three distinct prongs (Fig. 5). Opening of 
prothoracic gland visible. Coxae with spindle-shaped seta on cephalic, dorsal margin. D1 
of Al—A6 on reduced pinacula; D1 of A8, A10 on flat pinaculi. 

Fourth instar (n = 5). Length 13.1 mm (n = 8). Coloration similar to fifth instar. Head 
diam. 2.1 mm; height of frons 0.9 (0.8-1.1) mm; height of epicranial suture 1.1 (1.0-1.1) 
mm. Prothoracic scoli with reduced prongs (Fig. 6); adenosma well developed; spindle- 
shaped seta present on all thoracic coxae. D1 of all abdominal segments nearly indistin- 
guishable from secondary setae. Al0 prolegs completely reduced, with ring of setae 
surrounding undeveloped plantae. 

Fifth instar (n = 7). Length 34 (32-36) mm. Head brown; body green with white flecks; 
prothoracic prominences brown; a broad, reddish brown (concolorous green in some 
individuals) dorsal stripe extending from prothorax to tenth segment; a faint, horizontal, 
sublateral, white stripe from prothorax to tenth segment; thin, horizontal, midventral and 
subdorsal white stripes on thorax. Abdominal segments with diagonal white stripes ex- 
tending intersegmentally. Subventral and underside brown with white flecks. Head diam. 
3.3 mm; height of frons 0.9 (0.8-1.1) mm; height of epicranial suture 1.6 (0.5-1.8) mm. 
Mandibles as in Fig. 3. Hypopharyngeal complex as in Fig. 10; clypeus deeply invaginated 
with two pairs of stout setae on tip, and two pairs of minute setae on underside. Six larval 
stemmata; O83 surrounded by enlarged, sclerotized ring. Chaetotaxy as in Fig. 11. XD1 
of prothorax on a smooth, conical prominence (Fig. 7). Everted adenosma bifurcate (Fig. 
12). Spindle-shaped coxal setae as in Fig. 18. Abdominal chaetotaxy as in Fig. 9, two md 
microsetae on first abdominal segment. Prolegs on A8 with 30 (21-33) crochets, on A4 
with 34 (33-35) crochets, on A5 with 34 (32-37) crochets, on A6 with 35 (34-37) crochets. 

Pupa (n = 8). Length 18.2 (15.0-27.0) mm. Vertex with slight indentation. Tbl, Tb2, 
and Tar8 visible. Caudal edge of mesonotum with row of pits below ecdysial line of 
weakness. Spiracles with densely packed, short, recurved setae. Cremaster two oblique, 
outwardly pointing processes. 

Adult. Ocelli absent; antenna *% pectinate in both sexes, last 15 segments laminate, 
pectinations bicolored, black dorsally, ventral extension light brown. Epiphysis % length 
of tibia. One pair of tibial spurs on mesothoracic legs; spurs with grooved inner surface 
extending more than % spur length. Metathoracic tibia with two pairs of spurs; first pair 
with serrate ridges on inner tip surface approximately 4 spur length; second pair like 
mesothoracic one. All tarsi spinose; 5th tarsomere with 2 long, dorsolateral setae. Claws 
bifid. Male eighth sternite as in Fig. 14; male genitalia as in Figs. 15, 16. Female genitalia 
as in Fig. 17. Female wing with 5 frenular bristles. 


DISCUSSION 


Three chaetotaxic changes occur among instars in Litodonta hydro- 
meli. First, a spindle-shaped seta on the thoracic coxae appears in the 
third, fourth, and fifth instars. The function of this seta is unknown. It 
occurs in mature larvae of Furcula borealis (Guér.-Méneville), Het- 
erocampa astartioides Benjamin, Schizura unicornis (J. E. Smith), Hy- 
parpax aurora (J. E. Smith), and Disphragis sp.; but not in Dasylophia 
sp., Symmerista albifrons (J. E. Smith), Lochmaeus bilineata (Pack- 
ard), Hapigia sp., and Misogada unicolor (Packard). 

Other setal differences between first and subsequent instars of Li- 
todonta hydromeli concern secondary setae and length of SV1 on the 
first abdominal proleg. Scattered, short, secondary setae occur on the 
head and body in second and later instars. There are secondary setae 
on larval prolegs in all instars. In the first instar, SV1 on the first 


VOLUME 41, NUMBER 4 191] 





1] 
ne 


SS 





0.5mm 





Fics. 10-14. Litodonta hydromeli larval and adult morphology. 10, Hypopharyngeal 
complex of fifth instar; 11, Head chaetotaxy of fifth instar; 12, Everted adenosma of 
fifth instar; 13, Prothoracic leg of fifth instar; 14, Adult male eighth sternite. Scales 
represent 1 mm unless otherwise indicated. 


abdominal proleg is twice as long as other SV setae (Fig. 8). In all other 
instars, SV1 is the same length as other SV setae. 

Litodonta hydromeli pupae possess a character found in other North 
American Heterocampini. Mosher (1917) described the row of deep 
pits on the pupal thoracic dorsum in Schizura ipomoeae (Doubleday), 
S. concinna (J. E. Smith), Heterocampa guttivitta (Walker), and Loch- 
maeus bilineata (Packard). These pits also occur in Schizura unicornis 
(J. E. Smith), Disphragis sp. (Ecuador, culture No. WE84-007), Hy- 
parpax aurora (J. E. Smith), Heterocampa astartioides, and one species 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 15-17. Litodonta hydromeli genitalia. 15, Male genitalia except aedeagus; 16, 
Aedeagus; 17, Female genitalia. Figs. 15 and 16 are same scale. Scales represent 1 mm. 


of Nystaleini, Strophocerus punctulum (Schaus). The taxonomic dis- 
tribution of this character needs further study. 

Like Litodonta hydromeli, some larval Heterocampini possess elab- 
orate spines in the early instars (Packard 1895), and develop ventral, 
prothoracic defense glands which become functional in later stages 
(Herrick & Detwiler 1919, Eisner et al. 1972). Secretions of these glands 
in Schizura concinna and Lochmaeus manteo (Doubleday) contain a 
mixture of formic acid and straight-chain ketones (Eisner et al. 1972, 
Weatherston et al. 1979). These secretions deter invertebrates and ver- 
tebrates (Eisner et al. 1972). Notodontids are the only lepidopterans 
known to produce defense compounds containing ketones (Blum 1981). 

During larval development of Litodonta hydromeli, a progressive 
reduction in the dorsal armature occurs concurrently with the devel- 
opment of the ventral, prothoracic defense gland. The gland can be 


VOLUME 41, NUMBER 4 193 


dissected out of second and later instars. It is the same shape as that of 
Schizura concinna (Weatherston et al. 1979). The spray of fifth instars 
has a strong, acidic odor and deters Atta texana (Buckley) (Weller 
unpubl.). The ants vigorously clean their antennae after encountering 
a larva. However, the spray is not effective against another local ant, 
Camponotis sp. 

In addition to ontogeny of defense structures, ontogeny of feeding 
behavior occurs. Changes in feeding behavior during larval develop- 
ment are correlated with changes in mandibular morphology. First 
instars feed on the upper surface of the leaf and skeletonize it with 
forklike mandibles (Fig. 3), whereas subsequent instars ingest the entire 
leaf, and fifth instar mandibles have a thin, ridged, cutting edge and 
an inner, roughened surface (Fig. 4). Similar feeding behavior changes 
are known for several notodontid species (Godfrey & Appleby 1987). 


ACKNOWLEDGMENTS 


I thank E. Taylor for technical assistance. Daniel Janzen provided the pupa of Stro- 
phocerus punctulum. I also thank the following for comments on the manuscript: W. 
Lybarger, J. Miller, J. Rawlins, R. Robbins, M. Ryan, and an anonymous reviewer. Elaine 
Hodges gave extensive advice on the figures. This work was supported by Sigma Xi, and 
the research conducted at the Brackenridge Field Laboratory, Austin, Texas. 


LITERATURE CITED 


BLum, M.S. 1981. Chemical defenses of arthropods. Academic Press, New York. 562 pp. 

Dyar, H.G. 1904. Description of the larva Litodonta hydromeli Harvey. Proc. Entomol. 
Soc. Wash. 6:3-4. 

EISNER, T., A. F. KLUGE, J. C. CARREL & J. MEINWALD. 1972. Defense mechanisms 
of arthropods. XXXIV. Formic acid and acyclic ketones in the spray of a caterpillar. 
Ann. Entomol. Soc. Am. 65:765-766. 

ForBEs, W. T. M. 1948. Lepidoptera of New York and neighboring states. Part II. 
Geometridae, Sphingidae, Notodontidae, Lymantriidae. Cornell Univ. (New York) 
Agric. Exp. Sta. Ithaca Mem. 274. 268 pp. 

GopFREY, G. L. 1984. Notes on the larva of Cargida pyrrha (Notodontidae). J. Lepid. 
Soc. 38:88-91. 

GopFREY, G. L. & J. E. APPLEBY. 1987. Notodontidae (Noctuoidea). In Stehr, F. W. 
(ed.), Immature insects. Kendall-Hunt, Dubuque, Iowa. 754 pp. 

Harvey, L. F. 1876a. New Texan moths. Can. Entomol. 8:5-7. 

1876b. Notes on Litodonta, with remarks on Oncocnemis. Can. Entomol. 8: 
109-110. 

HERRICK, G. W. & J. D. DETWILER. 1919. Notes on the repugnatorial glands of certain 
notodontid caterpillars. Ann. Entomol. Soc. Am. 12:44—48. 

HINTON, H. 1946. On the homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol. 
Soc. London 97:1-87. 

HOLLAND, W. J. 1903. The moth book. Dover, New York. 479 pp. 

Ho.tioway, J. D. 1983. The moths of Borneo. Part 4. Notodontidae. Malay. Nat. J. 37: 
1-107. 

KIMBALL, C. P. 1965. Arthropods of Florida and neighboring land areas. Vol. 1. Lep- 
idoptera of Florida. Div. Plant Industry Florida Dept. Agric., Gainesville. 363 pp. 





194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


MOosHER, E. 1917. Pupae of some Maine species of Notodontoidea. Maine Agric. Exp. 
Sta. Bull. 259. 84 pp. 

PACKARD, A. S. 1895. Part 1. Natedtrtidas. Mem. Nat. Acad. Sci., VII, pp. 7-287. 

PETERSON, A. 1948. Larvae of insects. Part I. Lepidoptera and i infesting Hyme- 

noptera. Published by author, Columbus, Ohio: 315 pp. 

WEATHERSTON, J., J. E. PERCY, L. M. MACDONALD & J. A. MACDONALD. 1979. Mor- 
phology of the prothoracic defensive gland of Schizura concinna (J. E. Smith) (Lep- 
idoptera: Notodontidae) and the nature of its secretion. J. Chem. Ecol. 5:165-177. 


Received for publication 2 February 1987; accepted 11 August 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 195-198 


HOSTS, BIOLOGY, AND DISTRIBUTION OF 
ZALE PHAEOCAPNA (NOCTUIDAE) 


TIM L. MCCABE 


Biological Survey, New York State Museum, State Education Department, 
Albany, New York 12230 


ABSTRACT. First-instar Zale phaeocapna Franclemont (Noetuidae: Catacolinae), 
when offered hosts from the immediate environs of the parental female in the Adirondack 
Mountains, New York, fed on Corylus cornuta Marsh. and C. americana Walt. (Cory- 
laceae). Two field-collected larvae, compared to the bred larvae and judged conspecific, 
were beaten from Ostrya virginiana (Mill.) K. Koch (Corylaceae) in Florida. A second 
clutch of larvae was reared on Hamamelis virginiana L. (Hamamelidaceae). The larva 
is described, illustrated, and compared to Zale minerea (Gn). 


Additional key words: Catacolinae, larvae, Zale minerea, Corylaceae. 


During field studies near Indian Lake in the Adirondack Mountains 
of New York, I obtained a gravid female of Zale phaeocapna Francle- 
mont which oviposited in captivity. The resulting first instars were 
offered a selection of plants growing in the immediate environs. Corylus 
cornuta Marsh. (Corylaceae) proved to be an acceptable host. Acer 
rubrum L., Pinus strobus L., Abies balsamea (L.) Mill., Larix laricina 
(Du Roi) K. Koch, Alnus rugosa (Du Roi) Spreng., Betula papyrifera 
Marsh., Salix rigida Muhl., and Prunus virginiana L., the dominant 
trees at the site, were not accepted. I have since reared Zale phaeocapna 
repeatedly on Corylus cornuta and C. americana Walt. from ova of 
females taken on the Pine Bush Preserve in Albany Co., New York. In 
addition, I was sent two mature Zale larvae collected on Ostrya vir- 
giniana (Mill.) K. Koch (Corylaceae) in Torreya State Park, Florida, 
on 22 April 1979. I compared them to my bred Zale phaeocapna and 
determined them to be the same species. Another brood from the Pine 
Bush locale was switched in later instars to Hamamelis virginiana L. 
(Hamamelidaceae). Hamamelis forms buds later than Corylus, and its 
leaves are not present when first instars usually eclose. According to 
Hall (1952), the Corylaceae possibly were derived from an ancestral 
stock close to the present-day Hamamelidaceae. Certainly several noc- 
tuids can survive the shift from Corylus to Hamamelis (Pyreferra spp.). 
All the plants known to serve as larval hosts for Zale phaeocapna are 
closely related. 

Zale phaeocapna was once a rare insect, originally described from 
Pennsylvania and Alabama (Franclemont 1950), but is now known from 
northern New York to northern Florida, and is apparently becoming 
more common. The moth is abundant on the Pine Bush in Albany, but 
was not collected there despite intensive collecting in the 1880's on the 
barrens. The species will come to bait as well as light, so the advent of 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 





Fics. 1-5. Zale phaeocapna. 1, Living larva, Adirondack Mts., New York; 2, Adult; 
3, Setal map and pattern of head. P = posterior head seta, L = lateral head seta, A = 
anterior head seta, Af = adfrontal seta, F = frontal seta, Fa = frontal puncture, C = 
clypeal seta. 4, Oral face of left mandible; 5, Hypopharynx. 


the ultra-violet collecting light does not account for its apparent in- 
creased abundance. Baiting was a popular collecting practice in the 
1880’s and was used on the Pine Bush. The Albany Pine Bush is a sandy, 
pitch pine-scrub oak barrens, and Corylus is a common shrub. Ha- 
mamelis is uncommon and Ostrya is rare, whereas the moth is common. 
In the immediate locale where Zale phaeocapna was collected and 
reared in the Adirondacks, Ostrya and Hamamelis were absent. 

The parental female oviposited on 21 May 1980, and first stage larvae 


VOLUME 41, NUMBER 4 197 





Fics. 6-9. Zale phaeocapna. 6, Setal map of prothorax, lateral expanded view (from 
middorsal to midventral line); 7, Setal map of 1st abdominal segment, lateral view (ventral 
seta not visible); 8, Setal map of 8th abdominal segment, lateral view (ventral seta not 
visible). D = dorsal seta, XD = primary seta, SD = subdorsal seta, L = lateral seta, SV = 
subventral seta. 9, Ocellar map. 


eclosed 29 May. Fully mature larvae were obtained by 1 July 1980. 
Larvae were reared in clear plastic containers under ambient conditions. 
The pupae overwintered and were held in a refrigerator at 4°C until 
30 March. Adults began emerging 1 April 1981, suggesting overwin- 
tering as a pharate adult within the pupal shell (N = 17), a supposition 
supported by early spring emergence of the adults in nature. Voucher 
specimens are in the New York State Museum. 


Description of Mature Larva 
(Figs. 1, 3-9) 


Total length 36-40 mm (N = 10). Coloration (living material): head silver-gray with 
irregular dark markings. Body with an alternating blue-gray and yellow striped pattern 


198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


as follows: a narrow, yellow dorsal stripe; a broad, blue-gray subdorsal stripe; a broad, 
yellow supraspiracular stripe; a broad blue-gray spiracular stripe; a fine, narrow, yellow 
subspiracular stripe. 

Head (Fig. 3). Average head width 3.2 mm; epicranial suture 1.6 mm; height of frons 
0.75 mm (N = 10). Ocellar interspaces as in Fig. 9. 

Mouthparts. Hypopharyngeal complex with distal and proximolateral teeth as in Fig. 
5. Mandible with inner tooth and ridges as in Fig. 4. Spinneret extending beyond second 
segment of labial palpus. 

Thoracic segments (Fig. 6). Cervical shield weakly sclerotized; prothoracic spiracle 
averaging 0.33 mm high. 

Abdominal segments. Ab-1 (first abdominal segment) shown in Fig. 7. Crochets a 
uniordinal mesoseries of 13, 24, 31, 33, and 34 for segments 3, 4, 5, 6, and 10, respectively 
(rounded to nearest whole number). Ab-8 (Fig. 8) spiracle 0.833 mm high, remaining 
spiracles 0.24 mm high; spiracles black. D1 (first dorsal) setal bases on Ab-8 markedly 
protuberant; protuberance slightly higher than height of Ab-8 spiracle. 

Material examined. 10 specimens, 10 km E of Indian Lake, Hamilton Co., New York, 
lat. 43°45'30” long. 74°10'14”, elevation 555 m. 

Diagnosis. In the original description of Zale phaeocapna, Franclemont (1950) consid- 
ered it allied to Z. minerea (Gn.) based on adult morphology. Superficially, the adults 
look like Z. galbanata (Morrison), but can be distinguished by the lack of an adterminal 
bar below M3 on the forewing (Fig. 2). The larvae of Z. minerea (based on 10 specimens 
in the New York State Museum as well as Crumb’s [1956] description) compares to that 
of Z. phaeocapna as follows: Z. minerea has a much broader oral tooth on the mandible; 
hypopharyngeal complexes appear the same; Z. minerea head capsule maculation is much 
more extensive, reaching to the front of the head; Z. phaeocapna has alternating longi- 
tudinal blue-gray and yellow stripes and is narrow-bodied, whereas Z. minerea is brown 
and thick-bodied. Zale minerea has been recorded on rose and willow (Crumb 1956), 
birch and many other trees (Forbes 1954). 


ACKNOWLEDGMENTS 


Dale F. Schweitzer and H. D. Baggett collected the Florida larvae. I thank Schweitzer 
for loaning specimens from the Yale Peabody Museum. John G. Franclemont successfully 
reared Z. phaeocapna on Hamamelis from ova that I provided from the Albany Pine 
Barrens. Steven Teale, Carol Kuhn-Teale, and Edward Blakemore assisted in the Adi- 
rondack rearings. Alexandra Leff produced Figs. 3 and 5. I thank Charles Sheviak for 
our discussion concerning the relations of Hamamelis. Christopher Supkis made the black 
and white print from my color slide for Fig. 1. Brian Farrell kindly granted permission 
to collect on his property. I thank G. L. Godfrey and an anonymous reviewer for comments. 
Contribution number 527 of the New York State Science Service. 


LITERATURE CITED 


CruMB, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agric. Tech. Bull. 1135. 
306 pp. 

ForBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Cornell 
Univ. Agric. Exp. Sta. Mem. 329. 433 pp. 

FRANCLEMONT, J. G. 1950. Notes on some genera and species of eastern moths with 
description of new species (Lepidoptera: Phalaenidae). Bull. Brooklyn Entomol. Soc. 
45:144-155. 

HALL, J. W. 1952. The comparative anatomy and phylogeny of the Betulaceae. Bot. 
Gaz. 113:235-270. 


Received for publication 9 March 1987; accepted 30 July 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 199-208 


HOST SPECIFICITY AND BIOLOGY OF 
PROCHOERODES TRUXALIATA (GUENEE) (GEOMETRIDAE), 
A POTENTIAL BIOCONTROL AGENT FOR THE 
RANGELAND WEED BACCHARIS HALIMIFOLIA L. 

IN AUSTRALIA 


W. A. PALMER 


North American Field Station, Queensland Department of Lands, 
2714 Pecan Drive, Temple, Texas 76502 


AND 


J. W. TILDEN 
125 Cedar Lane, San Jose, California 95127 


ABSTRACT. Prochoerodes truxaliata is an ectophagous foliage feeder native to the 
western United States. It is multivoltine, with larvae found throughout the year on 
Baccharis pilularis, its only known host. Normal larval development occurred in the 
laboratory only on species of Baccharis, including B. halimifolia. Larvae also developed 
on the closely related Chrysothamnus nauseosus, but slow growth and high mortality 
suggest it is not a natural host. The insect is considered sufficiently stenophagous for 
introduction into Australia to control Baccharis halimifolia. 


Additional key words: _ biological control, Baccharis sarathroides, B. neglecta. 


The woody shrub Baccharis halimifolia L. (Asteraceae: Astereae: 
Baccharineae), an introduction from North America, is a serious weed 
in Queensland, Australia (Stanley & Ross 1986). The Queensland De- 
partment of Lands, through the Alan Fletcher Research Station, has 
instigated a long-range research program to find biological control agents 
in the New World for release against this weed in Australia. 

One source of potential biocontrol agents is the fauna feeding on 
species closely related to the weed. Indeed, some authors (Pimentel 
1968, Hokkanen & Pimentel 1984) suggest that such insects may be 
better biocontrol agents because they possess less “ecological homeo- 
stasis’. Programs against Opuntia spp. (Dodd 1929, Fullaway 1954, 
Pettey 1948) provide examples where insects from hosts other than the 
target species have given significant control. 

Baccharis pilularis is one of 20 United States species of the predom- 
inantly South American genus, and is found throughout coastal Cali- 
fornia and Oregon where two subspecies are recognized. Baccharis p. 
pilularis DC. is a prostrate form found near the coast and often grown 
as an ornamental. Baccharis p. consanguinea DC. is an erect form 
found further from the coast, and is morphologically very similar to B. 
halimifolia. 

Tilden’s (195la) comprehensive survey of the insect fauna associated 
with B. pilularis formed a useful adjunct to the biological control pro- 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


gram. A number of species listed there were studied further. One, 
Rhopalomyia californica Felt (Diptera: Cecidomyiidae), is already con- 
trolling B. halimifolia in some areas of Australia (McFadyen 1985). A 
second, Trirhabda flavolimbata (Mannerheim) (Coleoptera: Chryso- 
melidae), was found to have too wide a host range for introduction; 
laboratory tests indicated it might breed on Aster novae-angliae (Palm- 
er 1985). 

A third, Prochoerodes truxaliata (Guenée) (Geometridae), is the sub- 
ject of this paper, which reports studies conducted before permission 
was sought to introduce the insect for biocontrol purposes. Five of the 
seven species in the North American genus Prochoerodes are general 
feeders on trees and shrubs. The other two, P. truxaliata and P. am- 
plicineraria (Pearsall), are so distinct that they may not be congeneric 
with the others (D. C. Ferguson pers. comm.). Prochoerodes truxaliata 
has been recorded only from Baccharis pilularis, although adults have 
been collected in New Mexico, Utah, Colorado, and Arizona, all outside 
the range of B. pilularis. One specimen of Prochoerodes amplicineraria 
has been reared from Chrysothamnus nauseosus. 

Botanical nomenclature here follows Bailey and Bailey (1976) or 
Correll and Johnston (1979). 


BIOLOGY 


Moths (illustrated in Holland 1968) were nocturnal, emerging from 
resting places at dusk. They were observed hovering above Baccharis 
pilularis as late as 0100 h at Davis, California. Copulation occurred in 
early evening and oviposition commenced about three days after eclo- 
sion. Unmated females produced sterile eggs. One mated female laid 
231 eggs, and after death contained a further 119 undeveloped eggs, 
for a total brood of 350. Females oviposited on the plant, but as the 
eggs have little exochorion they usually fell to the ground. Females also 
oviposited as they crawled on the ground around the plant. 

Eggs were pale green, nearly spherical (0.8 mm diam.), smooth and 
glossy, but fertile eggs turned brown within 48 h. Eclosion occurred in 
8 to 10 days at room temperature (26°C). 

Larvae left eggs by eating a hole in the chorion without eating the 
shell, doubled their size by inhaling air, and exhibited negative geo- 
tropism by climbing any available object. The following nondiagnostic 
larval description is given only for special field use. First instars were 
dark gray with pale lateral vittae. The head was straw colored with 
numerous small dark dots, the absence of which from the frons gave 
the appearance of a frontal line. Thoracic legs and prolegs were also 
straw colored. Second instars were a translucent neutral gray with faint 


VOLUME 41, NUMBER 4 201 


dark lateral stripes, and lacked the disproportionately large head of the 
first instar. Third and later instars were brown, which intensified with 
maturity. Mature larvae were irregularly dotted with dull black, the 
dots coalescing to form poorly defined dorsal and lateral stripes. Sub- 
dorsal pale lines bordered the dorsal stripe, and two irregular light lines 
extended laterally, forming a margin to the lateral stripe. The first 
thoracic and abdominal spiracles were orange, often with blue centers. 
A pair of short fleshy protuberances was present on the fourth abdominal 
tergite, and there were indications of a second pair of protuberances 
on the fifth segment. Abdominal setae were black. In the laboratory, 
mean development times for 7 instars (N = 8) reared on B. pilularis 
were 4, 4, 5, 6, 7, 11 and 12 days, respectively. Two of 10 individuals 
underwent an 8th instar. 

Small larvae created “windows” in leaves but larger larvae consumed 
foliage from the leaf edge backwards, often consuming the whole leaf. 
Most larval growth and foliage consumption occurred in the last two 
instars. Fully grown larvae were 40-45 mm long. Larvae remained on 
the foliage during the day, but most feeding occurred at night. Resting 
larvae assumed a typical geometrid posture. Small larvae, when dis- 
turbed, dropped and returned on silk strands attached to the plant. 
Large larvae clung tightly to twigs and were difficult to dislodge. Mature 
larvae ceased feeding about three days before pupation and sometimes 
wandered away from the foliage. 

Pupation occurred in a slight cocoon made either in the foliage or 
on the ground. If on the plant, larvae gathered several leaves together 
with a few strands of silk to form the cocoon. Pupae were obtect, and 
15-20 mm long. Larvae were reared on clusters of Baccharis neglecta 
foliage to compare growth of the sexes. Male pupae were lighter (mean 
0.21 + SE 0.04 g, N = 90) than female pupae (0.32 + 0.07 g, N = 50). 
Development times from egg to adult averaged 66.2 + 0.5 (N = 90) 
and 70.0 + 0.6 (N = 50) days for males and females, respectively, at 
26°C. 

There were three generations a year at Davis and Stanford, Califor- 
nia, but larvae were present throughout the year, and overwintered in 
this stage. Large larvae were present at the end of winter, and produced 
moths in early April. A second generation was seen in mid-summer, 
and an autumn generation produced moths in October. 

Larvae were collected from both subspecies of B. pilularis, but in- 
frequently and then usually at low densities (one or two per bush). Very 
high densities (hundreds per large bush) were occasionally found near 
Davis, and these severely defoliated plants. The insect has occasionally 
become a pest of ornamental B. p. pilularis. 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


REARING 


A laboratory colony was started and maintained for a number of 
generations using the following procedure. Wild moths were collected 
and confined in paper cups for oviposition. Resultant neonate larvae 
were transferred to clusters of foliage (Baccharis pilularis, B. halimi- 
folia, or B. neglecta) held in a 37 X 27 x 17 cm plastic shoe box. 
Approximately 20 larvae were placed on each cluster. They were trans- 
ferred to fresh foliage twice a week. 

Pupae were collected from the container bottom and foliage, and 
placed with sugar-water wicks and branches of Baccharis in a plastic 
shoe box with the bottom replaced by fly mesh. Eggs fell through the 
fly mesh, and were collected from paper towelling under the cage. Eggs 
were refrigerated for about a month without ill effect; moths, larvae, 
and pupae could also be cooled for at least a week. 


Host SPECIFICITY 


Host specificity of P. truxaliata was determined by examining pinned 
specimens in major entomological collections, and conducting labora- 
tory trials to determine oviposition preference, neonate feeding, feeding 
behavior of late instars, and behavior in a multiple choice situation. 

Museum records. Collections at the University of California (Berke- 
ley, Davis, and Riverside); California Academy of Science, San Fran- 
cisco; Los Angeles County Museum of Natural History; National Mu- 
seum of Natural History, Washington, D.C.; and American Museum of 
Natural History, New York, were examined. Most specimens in these 
collections had been taken at light traps. Limited data on pinned spec- 
imens nominated B. pilularis as host. 

Oviposition preference. Cages approximately 1 m® were placed in a 
temperature-controlled glasshouse. Cages contained four potted plants, 
each of a different species, and each resting in a 30-cm diam. white 
dish in a cage corner. Sixteen laboratory-reared pupae (6 female, 10 
male) were placed in a cup at cage center with sugar-water wicks. After 
eclosion and oviposition, eggs in the white dishes were counted. The 
plants were further tended and examined every alternate day, and any 
larvae present on the foliage were counted. Each cage of four plant 
species was replicated twice. Egg numbers were analyzed by analysis 
of variance (using a log N + 1 transformation) to determine differences 
in ovipositional preference. Four such series using different plant species 
were studied and separately analyzed. 

Prochoerodes truxaliata exhibited an ovipositional preference for 
some species over others, with Baccharis halimifolia being a preferred 
host (Table 1). Larvae were found only on B. halimifolia and Chryso- 


VOLUME 41, NUMBER 4 203 


TABLE 1. Numbers of eggs collected from dishes surrounding potted plants after 
oviposition, and numbers of larvae observed on plants. Each series replicated twice. 


Mean maximum 


Mean no. eggs no. larvae 

Series 1 

Baccharis halimifolia L. (Tribe Astereae) 50 a* 25 

Chrysothamnus nauseosus (Pall.) Britt. (Astereae) 49 a 20 

Aster novae-angliae L. (Astereae) Ob 0 

Bellis perennis L. (Astereae) 2b 0 
Series 2 

B. halimifolia 89 a* 20 

Solidago altissima L. (Astereae) 40a 0 

Conyza canadensis (L.) Cronq. (Astereae) 19a 0 

Lactuca sativa L. (Lactuceae) 120a 0 
Series 3 

B. halimifolia 268 a* 3 

Chrysanthemum morifolium Ramat. (Anthemidae) 89 a 0 

Tagetes lucida Cav. (Tageteae) 89a 0 

Cynara scolymus L. (Cardueae) 389 a 0 
Series 4 

B. halimifolia 189 a* 13 

A. novae-angliae 18b 0 

Dahlia pinnata Cav. (Heliantheae) 3b 0 

Gaillardia pulcheila Foug. (Heliantheae) 4b 0 


* Means separated by different letters differ significantly (<0.05) by the LSD test from other means in the same series. 


thamnus nauseosus. Those on the former developed normally to pu- 
pation, but those on the latter developed much more slowly and with 
a very high mortality in later instars, only one larva successfully pu- 
pating and producing a normal looking moth. 

No-choice feeding of neonate larvae. Five unfed, neonate, laboratory 
reared larvae were placed in a paper cup with a young leaf of one 
plant species. Leaves were changed daily and after 72 h survival was 
assessed. Not all plants could be tested at the one time but Baccharis 
halimifolia was always included as a control with each series. Each 
treatment was replicated at least three times and, where possible, foliage 
was obtained from different plants for replication. 

Results (Table 2) show that larvae survived only on B. halimifolia, 
B. neglecta, B. pilularis, B. sarathroides, and Chrysothamnus nauseo- 
sus, and on these species survival was approximately 80%. Larvae did 
not survive on Baccharis glutinosa, B. bigalovii, B. pteronioides, or on 
22 other plant species. 

Feeding by later instars. The ability of late instars to develop on 
three plant species was evaluated. Larvae 20 mm long and approxi- 
mately 14 days old were selected from a colony raised on B. neglecta 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Survival of neonate larvae after 72 h exposure to leaves of various plant 
species. 


No. Mean percentage 
Plant replications surviva 
Tribe Astereae 
Baccharis halimifolia 1 80 
B. neglecta Britt. 1) 79 
B. pilularis DC. 6 96 
B. sarathroides Gray 3 80 
B. glutinosa Pers. 6 0 
B. bigelovii Gray 3 0 
B. pteronioides (DC.) Gray 3 ) 
Chrysothamnus nauseosus 8 92 
Isocoma wrightii (Gray) Rydb. 6 0 
Gutierrezia microcephala (DC.) Gray 8 0 
Aster novae-angliae 8 0 
Conyza canadensis 8 0 
Solidago altissima 3 0 
Tribe Anthemideae 
Leucanthemum maximum (Raymond) DC. 8 0 
Chrysanthemum morifolium 8 0 
Artemisia tridentata Nutt. 3 0 
Tribe Heliantheae 
Xanthium strumarium L. 3 0 
Parthenium hysterophorus L. 3 0 
Helianthus annuus L. 8 0 
Gaillardia pulchella Foug. 3 0 
Zinnia elegans Jacq. 8 0 
Dahlia pinnata Cav. 3 0 
Tribe Inuleae 
Antennaria fallax Greene 3 0 
Tribe Eupatoreae 
Eupatorium compositifolium Walt. 3 0 
Tribe Lactuceae 
Lactuca sativa L. 3 0 
Other families 
Vicia faba L. (Fabaceae) 8 0 
Lycopersicon esculentum L. (Solanaceae) 3 0 
Cucurbita pepo L. (Cucurbitaceae) 3 0 
Albizia julibrissan Durazz. (Mimoaceae) 3 0 
Delonix regia (Bojer) Raf. (Caesalpiniaceae) 8 0 


foliage. Five larvae were placed on foliage clusters of B. halimifolia, 
Chrysothamnus nauseosus (closely related to Baccharis, and on which 
neonate larvae were able to feed) and Aster novae-angliae L. (more 
distantly related to Baccharis, and on which neonates had not been able 
to feed). At intervals of 2 or 3 days for 14 days, larval lengths were 


205 


VOLUME 41, NUMBER 4 
LENGTH 
(mm.) 
36 +--+ B halimifolia ee 
ears. i 
BA @ C. nauseosus : 


H— - MK A novae-angliae 


TIME (Days) 


Growth in length of larvae on Baccharis halimifolia, Chrysothamnus nau- 





Bie. 


seosus, and Aster novae-angliae. Points represent means for 5 larvae. 
measured—a simple procedure because larvae naturally assume a stick- 


like posture on the leaves. 
Three distinctively different growth rates resulted (Fig. 1). While 
larvae survived for up to 12 days on Aster novae-angliae, fed somewhat 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Indices of preference in a multiple-choice experiment with two replications. 


Mean 
Mean percentage Mean no. 
distribution foliage leaves 
Plant index consumed nibbled 
Baccharis halimifolia (Tribe Astereae) 66 25 i 
B. neglecta 13 33 19 
Gutierrezia microcephala (Astereae) 2 <i 38 
Isocoma wrightii (Astereae) 18 <1 i 
Aster novae-angliae (Astereae) 0 0 0 
Xanthium strumarium (Heliantheae) <i 0 0 
Dahlia pinnata (Heliantheae) <I 0 0 
Zinnia elegans (Heliantheae) <] 0 0 
Leucanthemum maximum (Anthemideae) 0 0 0 
Chrysanthemum morifolium (Anthemideae) 0 0 0 
Tagetes lucida (Tageteae) 0 0 0 
Gerbera jamesonni Bolus ex Hook (Mutisieae) Il 0 0 


on the foliage and produced frass, negligible growth occurred, and after 
the first few days the larvae looked unhealthy. Those on Chrysothamnus 
nauseosus displayed an intermediate growth rate. 

Multiple-choice experiment. A multiple-choice experiment was con- 
ducted to determine whether larvae responded similarly on whole plants 
and cut foliage, and to observe their response when given a choice of 
plant species. A 53 x 69 x 84 cm cage covered with fly mesh was set 
up to contain 12 potted plants, each of a different species. Wooden 
planks were placed above the pots so that foliage and stems protruded 
through small holes in the planks which acted as a floor and facilitated 
larval movement between plants. Some hundreds of unfed neonate 
larvae and eggs were scattered over this floor, simulating the natural 
distribution of eggs on the ground around plants. The plants were 
examined daily for the duration of larval development, and the larvae 
on each plant counted. A distribution index (Palmer 1985) indicating 
relative larval abundance on each plant was calculated. Number of 
leaves attacked and an estimate of total feeding were recorded for each 
plant at the end of the experiment. 

Results (Table 3) indicated that Baccharis was the preferred host. 
Few larvae were seen on the non-Baccharis plants, and most of them 
were found in the first few days. None of the other species proved 
suitable, although some feeding was evident on the closely related 
Isocoma wrightii and Gutierrezia microcephala. Toward the end of 
the experiment, some of the Baccharis were considerably defoliated 
but there was no movement of the large larvae to other plants. 


VOLUME 41, NUMBER 4 207 


DISCUSSION 


Host specificity testing indicated that this insect is highly stenopha- 
gous, and that its host range is restricted to Baccharis species. Four 
Baccharis species appeared to be equally suitable hosts. This result 
suggests that B. sarathroides may be the host in Arizona, New Mexico, 
Colorado, and Utah where moths have been collected but B. pilularis 
is not found. A degree of affinity between Prochoerodes truxaliata and 
Chrysothamnus nauseosus was also evident, and reflects the phyloge- 
netic relation between Baccharis and other North American genera 
within tribe Astereae. This result, and the host range of another Cali- 
fornia insect, Aristotelia argentifera Busck (Gelechiidae), which is re- 
ported only from B. pilularis and Ericameria ericoides (Lessing) (Tilden 
1951b), support the hypothesis of B. J. Turner (pers. comm.) that Chrys- 
othamnus and Ericameria are the most closely related genera in North 
America to Baccharis. 

The insect seems sufficiently stenophagous for biological control use 
in Australia. It was also tested against a further 6 species in tribe Astereae 
and another 15 species in other tribes of Asteraceae including most of 
the commercially important ones. This degree of testing closely related 
species is greater than commonly done for potential biocontrol agents 
(Diatloff & Palmer 1987). The partial affinity with Chrysothamnus 
nauseosus is not important in the Australian context because this genus, 
like most North American genera of the tribe, is not found in Australia. 
Conyza canadensis, Solidago altissima and Aster novae-angliae are 
North American species introduced into Australia, and are probably 
the most closely related of the present Australian flora to Baccharis. As 
such, they might serve as the “critical test species’ advocated by Wap- 
shere (1975). Clearly, these plants were not suitable for Prochoerodes 
truxaliata. However, before final clearance for release in Australia, 
some testing against native species of Tribe Astereae should be under- 
taken there. 

Prochoerodes truxaliata might also be utilized as a biological control 
agent within the United States. In Texas, Baccharis neglecta and B. 
salicina, extremely closely related and often confused (Correll & John- 
ston 1979), are considered weedy, and have recently been recognized 
as a management problem on grazing lands (Scifres 1980). It might 
therefore be possible to introduce the insect into Texas to control these 
species. It is considered safe for introduction there (R. Bovey pers. 
comm.). 

Although ectophagous Lepidoptera have not been associated with 
many successful biological control programs, Prochoerodes truxaliata 
does have many desirable features of a good biocontrol agent. It is 


208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


multivoltine, highly fecund, capable of causing considerable damage 
to the host plant, and easily reared in the laboratory. Also, it does not 
have a strong diapause, a factor which might be particularly useful in 
Australia where winters are mild. There would appear to be good 
prospects of this moth establishing on Baccharis halimifolia if intro- 
duced for biological control. 


ACKNOWLEDGMENTS 


We thank L. E. Ehler, University of California, Davis, for help over the years; D. C. 
Ferguson for expert identification and helpful advice concerning Prochoerodes taxonomy; 
and B. L. Turner, University of Texas, Austin, for advice on phylogenetic relations of 
Baccharis to other genera. 


LITERATURE CITED 


BAILEY, L. H. & E. Z. BAILEY. 1976. Hortus third: A concise dictionary of plants 
cultivated in the United States and Canada. Macmillan, New York. 1290 pp. 

CORRELL, D. S. & M. C. JOHNSTON. 1979. Manual of the vascular plants of Texas. 
University of Texas, Dallas. 1881 pp. 

DIATLOFF, G. & W. A. PALMER. 1987. The host specificity of Neolasioptera lathami 
Gagné (Diptera: Cecidomyiidae) with notes on its biology and phenology. Proc. 
Entomol. Soc. Wash. 89:122-125. 

Dopp, A. P. 1929. The progress of biological control against prickly pear in Australia. 
Commonwealth Prickly Pear Board, Brisbane. 44 pp. 

FULLAWAY, D. T. 1954. Biological control of cactus in Hawaii. J. Econ. Entomol. 47: 
696-700. 

HOKKANEN, H. & D. PIMENTEL. 1984. New approach for selecting biological control 
agents. Can. Entomol. 116:1109-1121. 

HOLLAND, W. J. 1968. The moth book. A guide to the moths of North America. Dover, 
New York. 479 pp. 

MCFADYEN, P. J. 1985. Introduction of the gall fly Rhopalomyia californica from the 
USA into Australia for the control of the weed Baccharis halimifolia, pp. 779-787. 
In Delfosse, E. S. (ed.), Proc. VI Int. Symp. Biol. Contr. Weeds, August 1984, Van- 
couver, Canada. 

PALMER, W. A. 1985. The host range of Trirhabda flavolimbata (Mannerheim) (Co- 
leoptera: Chrysomelidae) and its suitability as a biological control agent for Baccharis 
spp. (Asteraceae: Astereae). Coleopt. Bull. 40:149-153. 

PETTEY, F. W. 1948. The biological control of prickly pears in South Africa. S. Afr. 
Dep. Agr. For. Sci. Bull. No. 271. 163 pp. 

PIMENTEL, D. 1963. Introducing parasites and predators to control native pests. Can. 
Entomol. 95:785-792. 

SCIFRES, C. J. 1980. Brush management. Principles and practices for Texas and the 
Southwest. Texas A&M University Press, College Station. 360 pp. 

STANLEY, T. D. & E. M. Ross. 1986. Flora of southeastern Queensland. Vol. 2. Queens- 
land Dep. of Primary Industries, Brisbane. Misc. Pub. QM84007. 623 pp. 

TILDEN, J. W. 195la. The insect associates of Baccharis pilularis De Candolle. Mi- 
croentomol. 16:149-188. 

1951b. Microlepidoptera associated with Baccharis pilularis. II. Tortricidae, 
Phaloniidae, Gelechiidae. Wasmann J. Biol. 9:239-254. 

WAPSHERE, A. J. 1975. A protocol for programmes for biological control of weeds. 
PANS 21:295-303. 





Received for publication 3 April 1987; accepted 9 September 1987. 


GENERAL NOTES 


Journal of the Lepidopterists’ Society 
41(4), 1987, 209-211 


TWO RELATED MIGRATIONS OF THE CALIFORNIA TORTOISE-SHELL 
BUTTERFLY IN MARIPOSA COUNTY, CALIFORNIA, IN 1986 


Additional key words: Nymphalidae, Nymphalis californica, hibernation. 


Mass exodus flights of the California tortoise-shell butterfly, Nymphalis californica 
(Boisduval) (Nymphalidae), usually follow defoliation of Ceanothus, the larval host, in 
years of abnormal temperatures and subnormal precipitation such as droughts (Shields 
1987, Utahensis 7(1):5-18). Years of peak migration in the Yosemite National Park region 
include 1911, 1922, 1933, 1961-62, 1971-72, and 1986-87. This note reports two related 
February—May 1986 migrations of N. californica near the SW end of Yosemite, and 
proposes the source to terminus route for the hibernant flight. 

In late July 1986 there were extensive migrations for 42 km in the Donner Pass region 
of Placer and Nevada counties; these flew SW and W (Knaus & Lambremont 1987, J. 
Lepid. Soc. 41:121-122), independent of the Yosemite migrations in source, timing, and 
(partially) direction. A syndrome of behavioral, physiological, and ecological character- — 
istics in migratory butterflies follows decreased production of juvenile hormone (Rankin 
1978, pp. 5-32 in Dingle (ed.), Evolution of insect migration and diapause, Springer- 
Verlag, New York; Herman & Dallmann 1981, J. Insect Physiol. 27:163-168). 

In 1986 at the Shields residence, Jerseydale, 1100 m elev., 14 km NE Mariposa, Mariposa 
Co., California, sightings were made along a NW-SE 80 m front in a pine forest clearing 
between cabins. Flight directions were recorded with a hand compass that had been 
checked against a surveyor’s compass. It was placed on level ground and oriviited to 
magnetic N (17'4°E of true N) to estimate flight directions. All times are PST. 

On 24 February 1986 in Skelton Canyon E of Jerseydale, worn hibernant N. californica 
were abundant (150-200 seen), most muddy areas having at least 10 or 20, sometimes 
40 or 50, in mid-morning on a warm, sunny, humid day (22°C). These were from a fall 
1985 emigration that had subsequently become resident. A small migration not partici- 
pated in by residents began on 26-27 February, and was confined to the canyon summit. 
A heavy rain (500 mm in Mariposa) fell on 15-19 February, followed during 24-28 
February by warm, clear, sunny, humid weather (low 20°’s C). 

This migration was first noticed on 28 February at Jerseydale (Table 1) beginning at 
0930 h and ceasing at 1605 h. Migrants flew up to 15-30 m above the ground, passing 
over pine trees instead of around them. A few changed direction when reaching the 
forest; these often appeared in groups of 2 to 4. A resident population (common but 
nonmigrating) was present at Jerseydale at this time. During March, no migration was 
seen on overcast days. A maximum migration of 75/h/30 m was reached on 1 March 
between 1100 and 1200 h (Table 1). After some rain and snow in early and middle March, 
the migration resumed during 19-27 March with 21-61/h/30 m, reducing to 1-2/h/30 
m from the end of March to 22 April. The flight rate (measured by car speedometer) 
was 32-40 km/h. In unforested areas between Darrah and Big Spring Hill, the flight was 
only 1-2 m above the ground. By late March many individuals had clear or tattered 
wings, and by mid-April many were exceedingly worn and flying slowly. In the early 
morning, the flight direction paralleled the incoming rays of sunlight. On 27 March, 
migrants were sometimes seen in 3's flying in V-formation. During its peak, migration 
began at 0920-0930 h and ceased at 1502-1605 h. In late February—early March, it was 
confined to an area between Jerseydale and Big Spring Hill (11 air km) with none migrating 
on either side of this front, such as at Briceburg or Mariposa. On 27 March a few N. 
californica were seen migrating ESE at 1100-1220 h on Hunter Valley Mtn., 610-915 
m elev., 29 air km NNW of Jerseydale. 


210 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. 1986 migrations of Nymphalis californica at Jerseydale. 


Percent in 
Time (h) Predominant Numbers/ predominant 
Date (PST) direction? 30-m front/h direction 
Hibernants 

27 February 1420-1500 SE oP 100 
28 February 13850-1415 SE 53> 100 
28 February 1420-1605 ESE 87° 100 

1 March 1100-1200 SE 75 90 
19 March 1250-1350 — Dp — 
21 March 1250-1350 SE 61 48 
22, March 1100-1200 SE 40 65 
24 March 1200-1300 ESE 26 46 
26 March 1200-1300 ESE 34 44 
26 March 1440-1540 ESE 24 al 
27 March 0920-1000 ESE 24> 34 
19 April 1100-1200 SE ”Y 100 

First Brood 

24 May 1230-1330 WNW 29 38 
25 May 0915-1015 E 12 33 
25 May 1430-1530 W 14 43 
25 May 1530-1630 W 5 80 
26 May 0815-0915 — 10 —4 
26 May 1130-1230 none 8 — 
27 May 1430-1530 W 8 38 
28 May 1024-1125 W 19 387 
28 May 1330-1430 NW 6 33 
31 May 1030-1130 W ii, WL 





«Predominant direction refers to greatest percent traveling in any one direction. 
> Rate/h unknown. 

¢ All but one flew SE to ESE. 

440% flew W and 40% flew E. 


The source area for the hibernant N. californica migration passing through Jerseydale 
was probably the Vaca Mts. W of Davis. Projecting the SE line of flight backwards 
intersects this region 230 km from Jerseydale (Fig. 1). There were hundreds to thousands 
of adults in canyons of the Vaca Mts. in February and March of the migration (A. Shapiro 
pers. comm.; News Lepid. Soc. 1987, no. 2, p. 17). By May there were fewer larval colonies 
in the Vacas than would have been expected from their earlier numbers. Projecting their 
SE line of flight forward from Jerseydale bisects the eastern Sierra Nevada between Lone 
Pine Creek and Big Pine Creek where D. Giuliani commonly saw nonmigrating adults 
from 26 February to 11 March of the migration. These were along streams in all canyons 
from 1980 m down to the valley floor (Fig. 1). None was seen there by 8-11 April 1986. 
This area is 185 km from Jerseydale and indicates a total of 420 km traveled by part of 
the migration. The distance between Lone Pine and Big Pine creeks is 65 km, indicating 
a 6-fold increase in width at the migration’s terminus when compared with the Jerseydale- 
Big Spring Hill width. The rate of travel from beginning to end was 1.5-2.4 days based 
on 420 km distance, 32-40 km/h flight speed, and 0920-0930 h to 1502-1605 h flight 
times. 

A brief migration of smaller, fresher, first-brood N. californica flew through Jerseydale 
later, in late May 1986 (5-29/h/30 m) (Table 1). On 20 May, 15 fresh adults were flying 
about, and 5 others were seen migrating. On 21-23 May, days were cool and overcast. 
The peak flight occurred on 24 May. Flight was near the ground and slower (10-12 km/ 
h) than the hibernant migration. On 26 May along Footman Ridge summit, 1400 m eley., 


VOLUME 41, NUMBER 4 ed ial 


»>2 





BIG PINE 
CREEK 


Fic. 1. SE migration of N. californica hibernants in 1986. 


1 km E of Jerseydale, 6 butterflies were seen migrating W to NW at 0955-1010 h. During 
late May, migration began at 0838 h and ceased at 1620 h, thus occupying a greater part 
of the day than the hibernant migration. For 1 h earlier and ¥% h later than this, specimens 
flew back and forth in the clearing and gully, basked, or alighted on trees. By 31 May 
most migrants were in intermediate wing-wear condition. Migration ceased during the 
first week of June. Three of 10 specimens collected on 24 May expelled meconia, indicating 
a nearby source area. This brood was likely the progeny of the hibernant migration. No 
N. californica were seen at Jerseydale during the summer until 25 August when one fresh 
specimen appeared. A few resident adults reappeared during the fall and early winter 
of 1986. 

Hibernant and first-brood migrations each had their own flight directions (SE vs. W). 
Greatest variability in direction occurred during breezy weather, with least variability 
on calm days. 

I thank L. P. Brower and two anonymous referees for comments on the manuscript. 


OAKLEY SHIELDS, 4890 Old Highway, Mariposa, California 95338. 


Received for publication 26 March 1987; accepted 29 July 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 212-213 


THE TYPE LOCALITY OF CATOCALA WHITNEYI AND 
REPORTS OF THIS SPECIES IN OHIO 


Additional key words: Noctuidae, Catocalinae, distribution. 


Catocala whitneyi Dodge has been reported from Ohio several times (Dyar, H. G., C. 
H. Fernald, G. D. Hulst & A. Busck 1902 [1903], Bull. U.S. Natl. Mus. 52, 723 pp.; 
Hampson, G. F. 1913, Catalogue of the Lepidoptera Phalaenae in the British Museum, 
Vol. 12, London, 626 pp.; Forbes, W. T. M. 1954, Lepidoptera of New York and neigh- 
boring states, Part III, Noctuidae, Cornell Univ. Agr. Exp. Sta. Mem. 329, Ithaca, New 
York, 433 pp.; Sargent, T. D. 1976, Legion of night: The underwing moths, Univ. of 
Massachusetts Press, Amherst, 222 pp.), but my efforts to locate valid Ohio specimens 
have been fruitless. After exhaustive investigation, I conclude that all previous references 
to the species occurring in Ohio are erroneous and explainable as misreading the type 
locality. 

G. M. Dodge (1874, Can. Entomol. 6:125-126) described C. whitneyi, giving the type 
locality as “Ohio, Ill.” This might be seen as a reference to the states of Ohio and Illinois, 
but in reality, Ohio is the name of a town in Bureau Co., Illinois. Dodge earlier (1874, 
Can. Entomol. 6:114-115) referred to Ohio, Illinois, as a single location. That Dodge was 
referring to this town is confirmed in a letter he wrote to James Angus: “In Bureau County, 
Ill., I found only Whitneyi”’ (Angus, J. 1884, Papilio 4:35-37). 

A credible source for the reports of C. whitneyi from Ohio is found in Hampson 
(above). Hampson lists five specimens and indicates that at least one type specimen is 
from Ohio: “Hab. U.S.A., Ohio, 1 4, 2 2 type, Illinois, Nebraska, Kansas (Snow), 1 4, 1 9.” 
This information contradicts W. Beutenmitiller (1907, Bull. Am. Mus. Nat. Hist. 23:145- 
151) who wrote, “The types of whitneyi were unfortunately destroyed by fire, as I am 
informed by Mr. Dodge, and the specimen in the Grote Collection in the British Museum, 
supposed to be the type is not one of the specimens from which the description was 
made.” Based on British Museum photographs of the five specimens listed by Hampson, 
including full label data, none of the specimens is from Ohio. The one specimen bearing 
a label with the word “Type” simply has the locality as “U.S. America, Grote Coll., 81- 
116.”° Two of the other specimens bear labels with data identical to that just cited and 
two specimens are labeled “Kansas, Snow, Grote Coll., 81-116.” 

Conceivably, C. whitneyi was collected by early Ohio lepidopterists. Charles Dury of 
Cincinnati and George Pilate of Dayton were accomplished lepidopterists and contem- 
poraries of Dodge. Grote identified many specimens for Dury, and Pilate provided the 
specimens for Grote’s description of C. dulciola (Grote, A. R. 1881, Papilio 1:5-6). The 
connection to Grote is important because one of the Grote specimens of C. whitneyi in 
the British Museum is allegedly a type. Either Dury or Pilate could have provided Grote 
with C. whitneyi, which would account for the Ohio record. This seems unlikely, however, 
since neither mentioned this species or the similar C. abbreviatella and C. nuptialis in 
their papers on Ohio Lepidoptera (Dury, C. 1876, Can. Entomol. 8:187-188; 1878, J. 
Cincinnati Soc. Nat. Hist. 1:12—-28; Pilate, G. R. 1882, Papilio 2:65-71). 

Ironically, W. Barnes and J. H. McDunnough (1918, Illustrations of the North American 
species of the genus Catocala, Mem. Am. Mus. Nat. Hist. 3(1), 47 pp.) must have known 
that C. whitneyi had not been recorded in Ohio when they described the distribution 
this way: “It has only been reported from a few of the Plains States from Nebraska and 
Kansas northward to southern Manitoba but appears to be fairly plentiful locally.” Had 
they explained the distribution in relation to the type locality, the enigma might have 
been resolved in 1918. 

The recorded foodplant of C. whitneyi, Amorpha fruticosa L. (Leguminosae), (Dodge, 
E. A. 1925, Entomol. News 36:267-268) occurs only infrequently along the Ohio River 
in western Ohio, but A. fruticosa is doubtfully native in Ohio (Braun, E. L. 1961, The 


VOLUME 41, NUMBER 4 213 


woody plants of Ohio, Hafner, New York, 362 pp.). Another probable foodplant, A. 
canescens Pursh, does not occur in Ohio. However, other species of Leguminosae are 
found in Ohio along with species of Catocala (minuta and illecta) that use them as hosts. 
Catocala whitneyi has been collected in Kentucky W of Louisville, where A. fruticosa 
is known to occur, but the moth has not been recorded from Indiana. 

No Ohio specimens have been located in Ohio collections, nor in major collections 
including the U.S. National Museum and the American Museum of Natural History. The 
Museum of Comparative Zoology, where remnants of the Pilate collection are housed, 
likewise has no Ohio specimens of C. whitneyi. 

The range of C. whitneyi reported by Barnes and McDunnough (above) is essentially 
correct. A specimen from Tennessee is in the Museum of Comparative Zoology. The 
range extends N and W through the Plains States including W Kentucky, Illinois, Wis- 
consin to Manitoba, and W through Missouri, Kansas, Nebraska, and N through the 
Dakotas. The recorded host, A. fruticosa, favors river and stream banks, whereas A. 
canescens is found on sandy soils and prairies. In Wisconsin (L. Ferge pers. comm.), C. 
whitneyi has a decided preference for prairie conditions, suggesting a relation with A. 
canescens; adults rest on the ground during the day. 

I thank R. W. Rings and T. D. Sargent for critically reviewing the manuscript, and C. 
W. Albrecht for encouragement. Many collectors and curators allowed access to collections 
under their care, and others, including C. V. Covell in Kentucky and E. M. Shull in 
Indiana, quickly responded to requests for data. I also thank F. H. Rindge and Mary 
Genett, American Museum of Natural History, for searching Beutenmiiller’s files, and 
the late Alan Hayes for specimen photographs and label data in the British Museum 
(Natural History). 


ERIC H. METZLER, Ohio Department of Natural Resources, Fountain Square C-2, 
Columbus, Ohio 43224. 


Received for publication 9 March 1987; accepted 13 August 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 213 


CORRECTION OF A NAME IN THE EPINOTIA VERTUMNANA 
(ZELLER) SPECIES-GROUP (TORTRICIDAE) 


Additional key words: taxonomy, Epinotia celtisana, Olethreutinae. 


In a revision of the Epinotia vertumnana (Zeller) species-group (Brown, R. L. 1986, 
J. Lepid. Soc. 40:327-346), Paedisca celtisana Riley (1881 [1882]) was transferred from 
synonymy with Paedisca vertumnana Zeller (1875) to synonymy with Proteopteryx 
laracana Kearfott (1907), with the latter name listed as the senior synonym. However, 
Epinotia celtisana (Riley) is the valid name by priority, and Epinotia laracana (Kearfott) 
becomes a junior synonym. 


RICHARD L. BROWN, Mississippi Entomological Museum, Drawer EM, Mississippi 
State, Mississippi 39762. 


Received for publication and accepted 11 August 1987. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 214-216 


TECHNICAL COMMENTS 


LOGIC AND PHYLOGENY: A CRITIQUE OF SCOTT’S PHYLOGENIES 
TO THE BUTTERFLIES AND MACROLEPIDOPTERA 


J. A. Scott (1985, J. Res. Lepid. 23:241-281; 1986, J. Res. Lepid. 25:30-88) proposed 
phylogenies to the Macrolepidoptera superfamilies (Fig. 1) and the butterfly families (Fig. 
8, in part). Although he presented an impressive amount of data on comparative mor- 
phology and behavior, in many cases these data do not support his phylogenies. Because 
nonsystematists might easily overlook this problem among the pages of morphological 
detail, I present one example from each paper showing that his data are inconsistent with 
his results. 


MACROLEPIDOPTERA 


Ability to hear the ultrasounds produced by bats and other predators evolved at least 
three times in moths (Sales, G. & D. Pye 1974, Ultrasonic communication by animals, 
Chapman & Hall, London, 281 pp.). The Geometroidea and Pyraloidea possess an ab- 
dominal tympanum, the Noctuoidea a thoracic tympanum, and the Choerocampinae 
(Sphingidae) a tympanum on the head (labial palps). Scott (1985, above) proposed that 
the Noctuoidea, Bombycoidea, Sphingoidea, and butterflies form a monophyletic group. 
His evidence was that the geometroid abdominal tympanum evolved into a thoracic 
tympanum in the ancestor of these taxa (point T in Fig. 1). As he stated, “The tympana 
moved to the metathorax.” 

The noctuoid-to-butterfly grouping is not supported by the data. The bombycoids, 
sphingoids, and butterflies lack the thoracic tympanum. Scott’s assumption that the ab- 
dominal geometroid tympanum is homologous with the thoracic noctuoid one is contra- 
dicted by the morphology and physiology of these structures (Forbes, W. T. M. 1916, 
Psyche 23:183-192; Richards, A. G. 19382, Entomol. Am. 13:1—48; Kiriakoff, S. G. 1952, 
Rev. Fr. Lepid. Fasc. 11-12:6 pp.; Maes, K. 1985, Nota Lepid. 8:341-350). No other 
characters support Scott’s noctuoid-to-butterfly grouping. A slightly different, but simpler 
phylogeny (Fig. 2) reflects the lack of support for the noctuoid-to-butterfly grouping and 
requires only one evolutionary change as opposed to two (gain and loss of the thoracic 
tympanum) in Fig. 1. 


BUTTERFLIES 


There are three major types of male forelegs among the butterflies (Bates, H. W. 1861, 
J. Entomol. 1:218-245; Ford, E. B. 1945, Butterflies, the new naturalist, Collins, London, 
368 pp.; Jander, U. 1966, Z. Tierpsychol. 23:799-844; Robbins, R. K. 1987, J. Lepid. Soc. 
40:138-157). 


TYPE I (Hesperiidae, Papilionidae, Pieridae). Foretarsi five-segmented with “spines,” 
sensilla, and pretarsal claws. Forelegs used for walking and cleaning the antennae. 


TYPE II (Lycaenidae sensu Eliot, J. N. 1973, Bull. Brit. Mus. (Nat. Hist.) Entomol. 28: 
371-505, including the Curetinae). Foretarsi fused into one segment, retain ‘spines’ 
and sensilla, but not pretarsal claws. Forelegs used for walking but not for cleaning the 
antennae. 


TYPE III (Riodininae, Libytheidae, Nymphalidae sensu Ehrlich, P. R. 1958, Univ. 
Kans. Sci. Bull. 39:305-370). Foretarsi partially or wholly fused, covered dorsally and 
ventrally with long scales (the “brush foot’’), devoid of “spines,” sensilla, and pretarsal 
claws, and greatly reduced in size. Forelegs not used for walking or cleaning the 
antennae. 


There are some exceptions to this summary (Type III forelegs occasionally have one or 
two ‘spines’ or sensilla, some male lycaenids have a segmented and clawed foretarsus), 
but they are irrelevant to my argument. 

Scott (1985, above) stated that the ancestor of the Lycaenidae-Libytheidae-Nymphalidae 


VOLUME 41, NUMBER 4 25 


ee ~ > a as & 
a OS OOS Ss OURO Or Ee 
A g » Wipacg> TK SS 
SG Saat eh OPQ I NL HERE 
VE 
L 
. 
1 2. 
Gis g wv 
es &% & ys S 
Se 
‘ S 
©) < O =e SS 
< Se Oar Se ne 
TW 
M 
Io 





3. 4. 


Fics. 1-4. 1, 2. Phylogeny to the Macrolepidoptera. The thoracic tympanum evolved 
at point T, but was lost at point L. 1, Scott’s phylogeny requiring two evolutionary steps; 
2, An alternate phylogeny requiring one change. 3, 4. Phylogeny to the butterfly “families.” 
Pupal midleg touching the eye evolved at point M. Evolution from one male foreleg type 
to another is represented by Roman numerals. The Styginae are omitted because of 
controversy over their male foreleg morphology (Forbes, W. T. M. 1960, Lepidoptera of 
New York and neighboring states, New York State College of Agriculture, Ithaca, 188 
pp.); 3, Scott’s phylogeny requiring five evolutionary steps; 4, An alternate phylogeny 
requiring three evolutionary steps. 


had a small leg that could not clean the antennae. Since Type III forelegs are the only 
ones that are significantly reduced in size, Scott’s statement implies that butterfly male 
forelegs evolved from Type I to Type III (in the lycaenid-nymphalid ancestor) to Type 
II (J-IJJ-II hypothesis). This hypothesis, however, is less parsimonious than a I-II-III 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


proposal in Bates (above). The I-II-III hypothesis requires foreleg walking to be lost once 
(change from Type IJ to III) while the I-JIJ-II hypothesis requires foreleg walking to be 
lost (change from I to III) and regained (change from III to I). 

Scott’s I-III-II hypothesis is inconsistent with his phylogeny (Fig. 3). The I-III-II hy- 
pothesis requires the Type II foreleg to evolve twice, once on the lineage to the Lycaenidae 
and once to the Curetinae (Fig. 3). The I-II-III hypothesis, on the other hand, implies an 

alternate phylogeny (Fig. 4) on which each male foreleg type evolves only once. 

Scott further supported his I-III-II hypothesis by noting that the pupae of Curetinae 
have the midleg touching the eye, as in Nymphalidae, but again, this information does 
not support his phylogeny. As background, the Curetinae possess a Type II male foreleg. 
Scott noted that the pupal midleg character state occurs in Curetinae, Libytheidae, and 
Nymphalidae, but it also occurs in Riodinidae (Chapman, T. A. 1895, Entomol. Rec. J. 
Var. 6:101-107, 125-131, 147-152). Scott’s phylogeny requires this character state to 
evolve twice (marked M in Fig. 3) while only one character change is necessary on the 
alternate phylogeny (point M in Fig. 4). 

Scott presented much information besides that on male forelegs, and his phylogeny 
(Fig. 3) may be better supported by these other characters than the alternate phylogeny 
(Fig. 4). The important point is not which phylogeny is “correct” but that Scott incorrectly 
supported his I-III-IJ hypothesis with male foreleg and pupal midleg characters. This 
finding casts doubt on the validity of his analyses in general. 

Phylogenies are basic to classification and to interpreting evolutionary hypotheses, but 
rigorously analyzed characters and character state distributions are needed to infer phy- 
logenies. Scott claims to use cladistic methods, but his analyses appear to be inconsistent 
with cladistic methodology (Lundberg, J. G. 1972, Sys. Zool. 21:398-413; Farris, J. S. 
1983, Adv. Cladistics 2:7-36). The prodigious amount of information that Scott presented 
on macrolepidopteran morphology and behavior will contribute to phylogenetic inference 
and, in this respect, is a major contribution to lepidopterology. However, it does not 
strongly support his conclusions. 

I gratefully acknowledge John Burns, Gerardo Lamas, Scott Miller, Michael Pogue, 
Alma Solis, and Susan Weller for reviewing this comment. 


ROBERT K. ROBBINS, Department of Entomology, NHB STOP 127, Smithsonian 
Institution, Washington, D.C. 20560. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 216-218 


LOGIC AND PHYLOGENY: REPLY TO R. K. ROBBINS 


Robbins is correct in questioning the homology of the noctuoid tympanum with other 
tympana. About the only use of tympana is to help indicate that Geometroidea split off 
the Macrolepidoptera line before Noctuoidea, although its detailed structure may provide 
useful traits within each superfamily. A fourth origin of the tympanum may be indicated 
by the dorsal as well as the usual ventral abdominal tympanum in Habrosyne (Thyatir- 
idae). Strong characters are used to devise branching schemes, and weak characters such 
as the tympanum are merely dragged along to wherever the strong characters place them. 
The position of Noctuoidea in J. A. Scott (1986, J. Res. Lepid. 25:30-38) merely minimizes 
the number of character changes in the overall Macrolepidoptera tree. Geometroidea and 
Noctuoidea seem the most primitive Macrolepidoptera because their larvae generally 
lack secondary setae and retain uniordinal crochets, their pupae retain the temporal 
cleavage line and the visible prothoracic femur, adults retain ocelli and the upper sector 
of the paracoxal sulcus, and, with Bombycoidea, adults retain the parepisternal rift and 
an areole. Geometroidea is at the base of the Macrolepidoptera tree because its abdominal 
tympanum may be phylogenetically related to the Pyraloidea abdominal tympanum, and 


VOLUME 41, NUMBER 4 Pad lie 


because its flat eggs are more primitive than upright Noctuoidea eggs. The position of 
Noctuoidea after Geometroidea is also assigned by default because the cluster Bomby- 
coidea-Sphingoidea butterflies share five derived traits (16-20 of Scott, above) which place 
this cluster on its own branch. Therefore, even when we discard the Noctuoidea tympanum 
because it evolved independently, Noctuoidea will have to stay put until new evidence 
to the contrary appears. The possible origin of Bombycoidea-Sphingoidea at the X of my 
Fig. 1 before Noctuoidea of Robbins’ fig. 1 is equivalent to moving Noctuoidea to between 
Sphingoidea and Hesperioidea on Robbins’ fig. 1, so I also was uncertain about the position 
of Noctuoidea. Currently, only these statements seem clear within Macrolepidoptera: 1) 
Geometroidea is the most primitive and Noctuoidea is next; 2) Bombycoidea and Sphin- 
goidea are closely related; and 3) Hesperioidea-Papilionoidea are on their own branch. 
What is needed are new characters, which readers will hopefully provide. 

Robbins’ fig. 2 is improbable because we know that in nearly all cases two rather than 
three species evolve at one time, so a three-branch split is improbable on a phylogeny. 
Even if a three-branch split occurred during species-level evolution, the subsequent great 
animal extinction rate (estimated at 99%) would make the survival of all three taxa to 
the present exceedingly unlikely. Some authors draw as many as half a dozen lines 
branching from one point, but this merely reflects their uncertainty. 

Among butterilies, the varying degrees of degeneration of the foreleg, especially the 
male foreleg, are weak traits that merely follow the strong traits when branching sequences 
are devised. I (below, pp. 256, 266) did not state that forelegs evolved from type I to III 
to II, only that the ancestor of Nymphalidae-Libytheidae-Lycaenidae had small forelegs, 
so that antennal cleaning by the middle leg evolved. Modifications of the foreleg such as 
tarsal fusion, claw loss, and scale elongation, or reversals of these states, apparently came 
later and proceeded differently in the various taxa. Libytheinae-Nymphalidae and Ly- 
caenidae contain many groups with small forelegs, and they both clean the antenna with 
the middle leg; the logic that a small foreleg forced a switch from foreleg to middle leg 
cleaning seems inescapable. But just how small the ancestral foreleg was is not clear. 
Robbins assumes that it was his type III. Libytheinae was the first lineage to evolve from 
the nymphalid line, and its male foreleg is about one-half normal size while the female 
foreleg is about two-thirds normal size; even such a minimal reduction, occurring mainly 
in one sex, would have been enough to cause a shift to the middle leg. Or, a fusion of 
tarsal segments or loss of tarsal claws could have eliminated the ability of the leg to curve 
over the antenna shaft, reducing its utility in cleaning and causing the shift. Or, could a 
mere reduction of body size to lycaenid dimensions, together with a less-than-linear 
reduction of antennal shaft thickness due to a need to retain shaft rigidity to support the 
club, have reduced the ability of the antenna to flex backward with a small enough radius 
tobe cleaned by the foreleg? If true, this ancestor would have a small foreleg in absolute 
dimensions, but a normal foreleg relative to the small middle and hind legs. One can 
classify the forelegs in various ways, many of which do not fit Robbins’ I-II-III system, 
which is too simple and unnatural. For instance, Riodininae and Curetinae both have the 
male foreleg coxa extending spinelike below the articulation with the trochanter, an odd 
trait that may show their phylogenetic relatedness (both share other traits cited by Scott, 
J. A. 1985, J. Res. Lepid. 23:241-281, including the middle leg touching the pupal eye, 
noticed in Riodininae by Chapman, T. A. 1895, Entomol. Rec. J. Var. 6:129). Many 
Lycaeninae have a segmented and clawed male tarsus (Eliot, J. N. 1978, Bull. Brit. Mus. 
[Nat. Hist.] Entomol. 28:373-505), contrary to Robbins’ type II; Eliot (pp. 394-3895) argues 
that some groups have reacquired segmented and clawed male forelegs. Nymphalidae 
also show varying degrees of modifications of the tarsus (Ehrlich, P. R. 1958, Univ. Kans. 
Sci. Bull. 39:305-379). It seems difficult to avoid the conclusion that there have been 
many independent modifications of male foreleg details, including reversals. Robbins is 
correct that a I-II-III sequence would be more parsimonious; however, parsimony of 
entire phylogenetic trees overrides parsimony within a single character, and trees forced 
to obey Robbins’ J-II-III sequence would require numerous added character changes in 
the tree because this sequence requires Nymphalinae to be evolved from the Lycaeninae- 
Curetinae ancestor. 

Robbins’ fig. 4 is impossible because of the massive number of shared derived traits of 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Lycaenidae (including Riodininae, Miletinae, Curetinae, Lycaeninae), some newly dis- 
covered on the first stage larva by D. M. Wright. Fully 28 strong shared derived traits 
now define Lycaenidae, and seven shared derived traits define Nymphalidae (including 
Libytheinae) (Scott above, and Scott, J. A. & D. M. Wright, Butterfly phylogeny and 
fossils, in Kudrna, Otakar (ed.), Butterflies of Europe, Vol. 2, Aula-Verlag, Wiesbaden, 
in press). Just as the principle of parsimony has its final judgment on entire trees rather 
than single characters, phylogenies must be based on numerous characters—the more the 
-better—and not on single characters. There will always be characters that are weak or 
difficult to interpret, or that show reversals, and even when worthless characters are 
discarded the remaining characters will not be of equal value; robust characters should 
be given greater weight. The shift to middle leg antenna cleaning is a strong character, 
but the detailed modifications of the male foreleg represent weak characters. The evo- 
lutionary history of weak characters is best determined by devising a phylogenetic tree 
using all characters (weighting the strong characters more heavily) and then using that 
tree to determine what happened to the weak traits. Using this method, the ancestral 
nymphalid-lycaenid foreleg may have shrunk to Libytheinae size, then later in Nym- 
phalidae the male and female foreleg shrank further, while in Lycaenidae the Libytheinae- 
type foreleg changed to a feather-duster type male foreleg in Riodininae, and in Ly- 
caeninae the foreleg became larger again, etc.; but all we know for certain is that the 
ancestral nymphalid-lycaenid foreleg was small in at least one sex. 

The character of the pupal middle leg touching the eye would have to evolve only 
once on my phylogeny (in the ancestor of Nymphalidae-Lycaenidae) contrary to Robbins, 
and would have to be lost only once (in the ancestor of Lycaeninae, because Riodininae 
also have the trait). 

It is good to question phylogenies, but one should not waste much time on weak 
characters; better to look for new characters, because the more one looks at a group of 
organisms, the more characters one finds. In most groups one can quintuple the known 
list of characters with hard work using morphology and behavior of all life stages. 

My two papers are ‘cladistic’ because they use the main two principles of cladistics, 
that a branch must be defined using shared derived traits, and that each branch must be 
monophyletic. Of numerous rules in cladistic variants, only those two rules are really 
necessary. It is also important to list all the character changes that must have occurred 
on the branches of the chosen tree to produce the character states observed in the living 
taxa; thus some weak characters are inevitably listed even though not given much weight 
in choosing the branching sequence of the chosen tree. 


JAMES A. SCOTT, 60 Estes Street, Lakewood, Colorado 80226. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 219-237 


OBITUARY 
ANDRE BLANCHARD (1896-1986) 


At 0730, 17 October 1986, the international lepidopterist community lost one of its 
most dedicated and capable amateurs. André Blanchard had been confined to his home 
at 3023 Underwood, Houston, Texas, for several years due to fibrosis of the lungs. Failing 
health made it necessary to discontinue in mid-November 1977 his most successful and 
rewarding field work on the Lepidoptera. Although confined, his interest in Lepidoptera 
did not diminish. Instead, he began studying in more detail the material at hand which 
resulted in the publication of 34 papers. In a personal communication dated 19 August 
1985 referring to a paper in preparation on Chlamydastis, he remarked: “This paper will 
almost certainly be my last”. It was. A year later on 20 August 1986 he fell in his home, 
fracturing a hip. He underwent surgery to repair the fracture, but after two months’ 
hospitalization the fracture did not heal, and due to general infection he passed away. 
His wife May Elise was at his bedside throughout the long hospital stay. Interment was 
in Calvary Cemetery, Houston, Texas, as he wished. 

André was born 1] June 1896 at Marennes (Charente Maritime), France, the youngest 
of two sons and a daughter (Suzanne) born to Francois Issac Blanchard and Marie Voyer. 
His father was affiliated with a banking facility. His older brother René was killed in 
action in World War I. In 1914 at age 18, through competitive examination, André was 
admitted to the Ecole Navale (French Naval Academy). Following graduation in 1917 
he served with the French Navy until 1929. 

While on active duty during World War I, André served on a battleship in the Med- 
iterranean, and a destroyer in the North Sea. During his last war year (1918), André was 
a seaplane pilot escorting ship convoys from Brest (Brittany) to Penzance (Cornwall). 
Early in 1919, following termination of the war, André was dispatched to New York to 
join the French battlecruiser Marseillaise. This marked his first visit to the New World 
which he found most enjoyable. In mid-1919 he returned to France and joined a group 
working under the leadership of Paul Langevin, a physicist doing research on the newly 
invented sonar. In April 1921, at his own request, André spent the next 12 months at 
Ecole Supérieure de Radiotechnique in Paris where he became a radio engineer. The 
remainder of his military career was at the Laboratoire du Centre d’Etudes de la Marine 
in Toulon, France. Here he not only taught radio but wrote the text which was later used 
at the Naval Academy. For two years (October 1927 to October 1929) he was commander 
of the 900-ton ship Les Eparges, especially equipped as the floating laboratory of the 
Centre d Etudes. 

Military honors received included the Croix de Guerre, 20 May 1917; Chevalier of the 
Legion of Honor, 8 January 1927; Officer of the Legion of Honor, 31 December 1950. 

In October 1929, Michelin Tire Company, Clermont-Ferrand, France, made André 
an offer which he could not refuse. He was placed in charge of the Company’s physics 
laboratory where he directed significant research on rubber, carbon black, heat-treatment 
of steel, and steel wire drawing. At the outbreak of World War II, André became head 
of the department that produced the new “metallic tire’. Thinking the patents might be 
sold to American tire companies in connection with the war effort, André was one of a 
delegation of five who left France in January 1941 for the United States to present the 
Michelin proposal. The deal failed and the delegation was denied exit visas until the end 
of the war. Not willing to remain inactive due to circumstances beyond his control, André 
went to work for the United States War Department translating from English to French 
military manuals ranging from Aerial Celestial Navigation to Optical Rangefinders. Mean- 
time in France, his wife and two daughters were being well cared for by Michelin. In 
October 1943, Schlumberger Well Services became aware of André’s availability, and 
hired him for the duration. After the armistice in Europe, André returned to France, his 
family, and Michelin. However, an overriding bond had been developed between André 
and Schlumberger; as a result, he returned to the United States in August 1946 with his 
family. Here he was again employed by Schlumberger, first as Manager of Engineering, 
later as Vice President of Engineering, and finally as Vice President of Research and 


220 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 





André Blanchard 


Development. He retired 30 June 1961 at age 65, but remained a paid consultant with 
Schlumberger for four more years. 

André’s family life began 22 April 1921 when he married Marguerite Normand at 
Loches, France. Born to this marriage were two daughters, Simone Blanchard Gerard, 
and Michelle Blanchard Gaymard. In 1958, following a long illness, Marguerite died, and 
on 9 May 1959 André and May Elise Moraud of Houston were married. No children 
were born to this union. André is survived by his devoted wife, May Elise, two daughters 
(Simone and Michelle), eight grandchildren and eight great-grandchildren. 

André’s avocation as a lepidopterist began in about 1925 when his wife, Marguerite, 
suggested that they should have a hobby. After thoughtful consideration they chose 
butterfly collecting. To house the European butterflies they would catch, some 30 glass- 
top specimen cases with cork pinning bottoms were manufactured. Collections were made 
as time permitted during the remaining years they spent in Europe. Unfortunately, there 


VOLUME 41, NUMBER 4 papay | 


was little time in his busy schedule to properly curate the collection, and it was virtually 
destroyed by dermestids. Nevertheless, when André and his family moved to the United 
States, the specimen cases came along and served as initial storage for his collection of 
Texas specimens. 

André joined the Lepidopterists’ Society in 1957. His primary interest at that time was 
butterflies. It was a paper of mine in 1959 (J. Lepid. Soc. 13:221-228) that brought us 
together. On 30 October 1960 he wrote, “I have been myself hoping for two to three 
years to find the foodplant of Phyciodes texana. I have not been as lucky as you but I 
feel very definitely that one more plant should be added to your list of four—Dicliptera 
brachiata.”” On 26 November 1960 we met and together in Memorial Park (Houston, 
Texas) learned that indeed Dicliptera brachiata is used in nature by this species. That 
meeting marked the beginning of a most enjoyable friendship between our two families 
for more than 25 years. During one of our early meetings, André remarked: “There is 
no need for both of us to work on the butterflies of Texas, I'm going to work on the 
moths.” 

Upon retirement André began collecting both seriously and enthusiastically. At first 
he used bait traps, UV and mercury-vapor lights set on a white bedsheet. In 1962 the 
late Perry A. Glick showed André a light trap used by the U.S. Department of Agriculture 
for sampling economic nocturnal insects. On 24 February 1963 he visited Joe P. Hol- 
lingsworth, College Station, Texas, who was working on a new model light trap for the 
Department. Measurements were taken and a sketch made of the new model; four days 
later André had his light trap. After putting the trap into use he learned that during 
certain times of the season many Coleoptera would come to the light, and because beetles 
are so slow to die they would virtually destroy Lepidoptera in the trap. This problem 
was essentially solved by a novel modification he made which separated most Coleoptera 
from Lepidoptera. 

Now, using three modified light traps, André started sampling nocturnal Lepidoptera 
statewide. Field trips were planned following the new moon which provided the longest 
periods of darkness. During the 16 years that followed many field trips were made 
throughout Texas resulting in the first statewide survey of noneconomic Lepidoptera 
(moths). May Elise accompanied him on every field trip, helped set up the traps in the 
evening, and helped pick them up early the following morning. After sorting the catch, 
specimens to be kept were placed in a relaxer, later pinned and spread in the field. Spread 
specimens were placed immediately in a dryer. Thus, except for the last catch of the 
trip, specimens taken were ready to be labeled and placed in the collection upon returning 
home. Many of the unwanted specimens were papered by May Elise to be given to 
correspondents or to museums. 

In early 1962 André was already thinking of preparing a checklist for Texas moths. 
To this end, a card file of species recorded from the State was initiated. At the same time 
additional specimen cases were being made to house the eventual collection of some 
60,000 spread specimens. He also began learning the technique of dissecting genitalia. A 
special collection of literature was accumulated. He was already expert at photography. 

Almost from the beginning of his intensive work on the moths of Texas, it became 
apparent that certain families and genera were in need of revision. As undescribed species 
accumulated this situation became a problem. He then began seeking help and advice 
from professional lepidopterists. Both he and his work were greatly respected within 
professional circles, thus the help needed came easy, although much too slow at times to 
please him. With the publication in 1983 of the Check List of the Lepidoptera of America 
North of Mexico, the way was cleared for at least a tentative State list but it came too 
late. Failing health demanded that he limit his activities to describing some of the new 
species he had collected. 

On 15 August 1977, André informed me that Edward C. Knudson, M.D., another 
amateur Texas lepidopterist, had visited him the day before, bringing along a box of 
moths for determination. He had been greatly impressed both by Knudson and the 
specimens brought for examination. As it turned out, this was the beginning of a continuing 
and deepening friendship between them. They worked together perfectly, and their 
manuscript production speaks for itself. Knudson will continue the statewide survey; he 
has prepared a tentative checklist representing more than 3000 moth taxa. 


222 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


André’s lepidopterological affiliations included: American Museum of Natural History 
(life member); The Lepidopterists’ Society, regular and sustaining member (1957-86), 
member-at-large of the Executive Council (1971-73), president (1975-his presidential 
address, J. Lepid. Soc. 30:1-4); The Lepidoptera Research Foundation (charter and regular 
member to July 1985); The Entomological Society of Washington (regular member 1968- 
86); Southern Lepidopterists’ Society (regular member 1985-86) from which he received 
the John Abbot Award 19 October 1985 for significant research on the Lepidoptera of 
Texas. 

Currently, fellow lepidopterists have described eighteen species honoring André and 
May Elise. Doubtless others will follow. A list of these patronyms together with other 
pertinent data is given in a later section. 

André prearranged disposition of his specimens, and “tools’’ for collecting and studying 
Lepidoptera. His primary collection of Texas insects went to the National Museum of 
Natural History (USNM), Smithsonian Institution, Washington, D.C. Based on an inven- 
tory by that institution (J. Lepid. Soc. 39:235—236), the collection totaled 76,852 specimens 
of which 16,305 were Coleoptera and 60,233 Lepidoptera. The latter, largest single 
accumulation of Lepidoptera ever made from Texas, included 82 holotypes, 700 paratypes, 
and 4600 microslides, mostly of genitalia. Both micro- and macro-Lepidoptera are rep- 
resented with more than 65 percent in four family groups: Noctuidae 18,500+, Pyraloidea 
9000+, Geometridae ca. 8000 and Tortricoidea ca. 4000 specimens. Frederick H. Rindge, 
American Museum of Natural History (AMNH), New York, New York, advised (21 
November 1986) that from 1958 through 1976 the AMNH received from Blanchard 
16,500 specimens of Texas Lepidoptera, and before this, about 1943, a number of spec- 
imens from the islands,of Martinique and Guadaloupe were donated by him. Of course 
many other institutions and individuals received small numbers of specimens, including 
paratypes, which André had collected or described. 

André’s “tools” for studying Lepidoptera included library material, optical equipment, 
photographic equipment, a card file of species taken or recorded from Texas, correspondence 
dealing with Lepidoptera, specially designed spreading boards, UV light traps, and spec- 
imen cases. The library material was given to Rice University, Edward C. Knudson, and 
Roy O. Kendall; the optical equipment and spreading boards to Knudson and Kendall; 
specimen cases (31) brought from France, to Kendall, other cases to USNM with the 
primary collection; all remaining “tools” to Knudson. 

Finally, on behalf of André’s friends, I paraphrase in part a classical eulogy: Rest, dear 
friend, lie down for an aeon or two until the Master of all good Lepidopterists shall put 
you to collecting anew. You will long be remembered by those of us who knew you, and 
your scientific contributions to the advancement of lepidopterology will live forever, but 
we think you knew that. 


PATRONYMS 


Lepidopterists have described eighteen species honoring André and May Elise Blan- 
chard. It seems appropriate that the list include those named for May Elise as well as 
André because they worked together in the field as a team. Still other patronyms honoring 
Blanchard will probably appear. The list of patronyms should be seen as a tribute to 
Blanchard for his significant contribution to the scientific advancement of lepidopterology, 
especially in Texas. No other individual has contributed so much toward a better un- 
derstanding of the Lepidoptera of Texas. 


Gelechiidae 
Dichomeris blanchardorum R. W. Hodges 1986. Gelechioidea, Gelechiidae (in part), in 
Dominick, R. B., et al., The Moths of America North of Mexico, Fasc. 7.1:48. Type 
Locality (TL): Texas, Cameron Co., Laguna Atascosa [National Wildlife Refuge]. 
Holotype 6 in USNM. 


Geometridae 


Plataea blanchardaria E. C. Knudson 1986. Proc. Entomol. Soc. Wash. 88:351-353. TL: 
Texas, Duval Co., 13 km W of Premont. Holotype ¢, 30 June 1985, in USNM. 


VOLUME 41, NUMBER 4 rages) 


Sicyopsis blanchardata D. C. Ferguson 1983. J. Lepid. Soc. 37:24-28. TL: Texas, Culberson 
Co., Guadalupe Mountains [National Park], Smith Canyon, 1753 m. Holotype 4, 22 
May 1973, in USNM. 

Stenoporpia blanchardi F. H. Rindge 1968. Bull. Am. Mus. Nat. Hist. 140:65-134 (116, 
117). TL: Texas, Brewster Co., Big Bend National Park, Basin. Holotype 4, 9 April 
1967, allotype-topotype 2, 4 October 1967, both in AMNH. 


Glyphipterigidae 
Drymoana blanchardi J. B. Heppner 1985. The Sedge Moths of North America (Hand- 
book No. 1). Flora & Fauna Publ., Gainesville, Florida, x + 254 pp. (49-53). TL: 
Texas, Jackson Co., Deutschburg [nr. Carancahua Creek, S of Francitas]. Holotype 
8, 7 October 1974, in USNM. 


Lasiocampidae 


Apotolype blanchardi J. G. Franclemont 1973. Mimallonoidea, Bombycoidea (in part), 
in Dominick, R. B., et al., The Moths of America North of Mexico, Fasc. 20.1:48, 
49. TL: Texas, Cameron Co., Brownsville. Holotype 6, 7 November 1969, in USNM. 

Tolype mayelisae J. G. Franclemont 1973. Mimallonoidea, Bombycoidea (in part), in 
Dominick, R. B., et al., The Moths of America North of Mexico, Fasc. 20.1:37, 38. 
TL: Texas, Brewster Co., Alpine. Holotype 2, 2 October 1963, in USNM. 


Noctuidae 


Basilodes [Stiria] blanchardi C. L. Hogue 1966. J. Res. Lepid. 4:275-280. TL: New Mexico, 
Eddy Co., Carlsbad Caverns National Park. Holotype 6, 17 September 1963, in Los 
Angeles County Museum of Natural History. 

Grotella blanchardi R. R. McElvare 1966. J. Lepid. Soc. 20:91, 92. TL: New Mexico, 
Eddy Co., White City. Holotype 4, 17 September 1963, in USNM. 

Opsigalea blanchardi E. L. Todd 1966. Proc. Entomol. Soc. Wash. 68:149-151. TL: Texas, 
Brewster Co., Alpine. Holotype 6, 9 September 1963, in USNM. 


Pyralidae 


Acrebasis blanchardorum H. H. Neunzig 1973. Proc. Entomol. Soc. Wash. 75:165-169. 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area, 1829 m. Ho- 
lotype 6, 5 June 1969, in USNM (type no. 72179). 

Homosassa blanchardi J. C. Shaffer 1976. Proc. Entomol. Soc. Wash. 78:431—434. TL: 
Texas, Harris Co., Houston. Holotype 6, 5 June 1967, in USNM (type no. 72179). 

Mojaviodes blanchardae E. G. Munroe 1972. Pyraloidea (in part), in Dominick, R. B., 
et al., The Moths of America North of Mexico, Fasc. 13.1B:188. TL: Texas, Presidio 
Co., Shafter. Holotype 3, 9 September 1969, in USNM. 

Pyrausta andrei E. G. Munroe 1976. Pyraloidea (in part), in Dominick, R. B., et al., The 
Moths of America North of Mexico, Fasc. 13.2B:127, 128. TL: Texas, Brewster Co., 
Big Bend National Park, Green Gulch. Holotype 2, 28 March 1971, in USNM. 

Scoparia blanchardi E. G. Munroe 1972. Pyraloidea (in part), in Dominick, R. B., et al., 
The Moths of America North of Mexico, Fasc. 13.1A:39, 40. TL: Texas, Jeff Davis 
Co., Davis Mountains, Mount Locke. Holotype 6, 6 September 1969, in USNM. 


Saturniidae 


Sphingicampa blanchardi D. C. Ferguson 1971. Bombycoidea (Saturniidae), in Dominick, 
R. B., et al., The Moths of America North of Mexico, Fasc. 20.2:47-50. TL: Texas, 
Cameron Co., Brownsville, Esperanza Ranch [now city housing]. Holotype ¢, no date, 
in USNM (type no. 71494). 


Sphingidae 


Amplypterus [Adhemarius] blanchardorum R. W. Hodges 1985. Proc. Entomol. Soc. 
Wash. 87:323-328. TL: Texas, Brewster Co., Big Bend National Park, Chisos Moun- 
tains, Panther Pass, 1829 m. Holotype 4, 4 June 1973, in USNM. 


224 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Tortricidae 


Rhyacionia blanchardi W. E. Miller 1978 in Powell, J. A. & W. E. Miller, U. S. Dept. 
Agric. Handbook 514, 51 pp. (19-21). TL: Texas, Montgomery Co., Conroe. Holotype 
6, 10 March 1968, allotype-topotype 2, 9 March 1971, both in USNM. 


BIBLIOGRAPHY 


At the time of his death two additional papers had been drafted in which Blanchard 
was junior author: 1) With R. W. Poole, describing two new genera and two new species 
of Noctuidae from Texas; and with J. G. Franclemont on the noctuid genus Protoperigea. 

The bibliography is divided into two sections, one dealing with European Lepidoptera 
(I-VIII), the other with North American Lepidoptera (1-59). Species treated in the latter 
are indexed under two headings: New Genera and Species, and Rare and Interesting 
Records. In each instance the new genera and species are arranged alphabetically and 
keyed to the numbered paper in the bibliography. 


Early Publications on Lepidoptera of France 


I. 1937. Thais rumina dans la vallée de la Jonte. L’Amateur de Papillons 8:252. 

II. 1937. Parnassius mnemosyne dans le Puy-de-Doéme. L’Amateur de Papillons 8: 
252, 253. 

III. 1938. Un arc-en-ciel d’ecailles de papillons. Rev. Frang. Lépid. 9:11-14. 

IV. 1938. L’Elevage de Macrothylacea rubi Linné. Rev. Frang. Lépid. 9:57-61. 

V. 1938. Capture en Auvergne d Heodes amphidamas Esp. (Lycaenidae). Rev. Franc. 
Lépid. 9:147-148. 

VI. 1940. Contribution a la connaissance de la faune des Lépidoptéres du Puy-de- 
Dome. Rev. Sci. Nat. d Auvergne 6:18-20. 

VII. 1940. Faune des Lepidopteres d’ Auvergne. Rev. Sci. Nat. d Auvergne (N.S.) 6(1- 
2):18-20. 

VIII. 1940. Observations sur quelques lépidoptéres d'Auvergne. Rev. Sci. Nat. d’Au- 
vergne (N.S.) 6(3-4):81-88. 


Publications on Lepidoptera of North America 


1. 1963. Contribution to the life history of Schoenobius maximellus (Pyralididae). J. 
Lepid. Soc. 17:234-235. 

2. 1964. The food plants of Syssphinx heiligbrodti (Saturniidae) in Texas. J. Lepid. 
Soc. 18:42. 

3. 1966. A new species of Glaucina (Geometridae) from Texas. J. Lepid. Soc. 22:247- 
250. 

4. 1968. New moths from Texas (Noctuidae, Tortricidae). J. Lepid. Soc. 22:133-145. 

5. 1969. A gynandromorphic Phaeoura mexicanaria (Geometridae). J. Lepid. Soc. 23: 
274-275. 

6. 1970. Observations on some Phycitinae (Pyralidae) of Texas with descriptions of 
two new species. J. Lepid. Soc. 24:249-255. 

7. 1971. Notes on three species of Heterocampa Doubleday with description of a new 
species. (Lepidoptera: Notodontidae). Proc. Entomol. Soc. Wash. 73:249-254. 

8. 1971. A new species in the genus Ursia Barnes & McDunnough (Lepidoptera: 
Notodontidae). Proc. Entomol. Soc. Wash. 73: 303-305. 

9. 1972. More new moths from Texas (Noctuidae). J. Lepid. Soc. 26:56-63. 

10. 1972. A new species of the genus Pyromorpha Herrich-Schaeffer (Pyromorphidae). 
J. Lepid. Soc. 26:79-82. 

11. 1973. Record and illustration of some interesting moths flying in Texas (Sphingidae, 
Ctenuchidae, Noctuidae, Notodontidae, Geometridae, Pyralidae, Cossidae). J. Lepid. Soc. 
27:103-109. 

12. 1973. A new species of the genus Glenoides McDunnough (Geometridae). J. Lepid. 
Soc. 27:141-1438. 

13. 1973. Two new species of Phycitinae from Texas, with description of two new 
genera (Pyralidae). J. Lepid. Soc. 27:219-225. 


VOLUME 41, NUMBER 4 225 


14. 1973. Erratum. J. Lepid. Soc. 27:278. (Corrects a name in J. Lepid. Soc. 24:249- 
255; Dioryctria auranticella (Grote) should be Dioryctria rossi Munroe.) 

15. 1975. A new phycitine genus and species (Pyraloidea). J. Lepid. Soc. 29:95-97. 

16. 1975. A new schoenobine genus and species (Pyraloidea). J. Lepid. Soc. 29:98- 
101. 

17. Rostrolaetilia—A new North American genus of the subfamily Phycitinae, with 
descriptions of seven new species (Pyralidae). J. Lepid. Soc. 29:131-150. 

18. 1976. Presidential address 1975—To my fellow amateurs. J. Lepid. Soc. 30:1—4. 

19. 1976. The genus Copablepharon in Texas, with description of three new species 
(Noctuidae). J. Lepid. Soc. 30:116-120. 

20. 1976. A new species of the genus Bertella Barnes & McDunnough (Pyralidae). J. 
Lepid. Soc. 30:211-213. 

21. 1976. Two new species of phycitine moths with description of a new genus (Pyrali- 
dae). J. Lepid. Soc. 30:284—288. 

22. 1976. Oenobotys texanalis. (With Eugene Munroe sr. author.) Pyraloidea (in part), 
in Dominick, R. B., et al., The Moths of America North of Mexico, Fasc. 13.2A:18-19. 

23. 1978. Atopothoures A. Blanchard a synonym of Goya Ragonot (Pyralidae). J. 
Lepid. Soc. 32:55-56. 

24. 1978. The status of Ollia parvella Dyar: Redescription in a new genus (Pyralidae). 
J. Lepid. Soc. 32:103-106. 

25. 1979. New status for Epiblema minutana (Kearfott) and new species of Epiblema 
Hiibner and Sonia Heinrich (Tortricidae). J. Lepid. Soc. 33:179-188. 

26. 1980. Five new species of the tribe Eucosmini (Tortricidae). J. Lepid. Soc. 33:209- 
215. 

27. 1981. A new species of the genus Peoria Ragonot (Pyralidae). J. Lepid. Soc. 34: 
338-339. 

28. 1981. Charadra ingenus Smith in West Texas (Lepidoptera: Noctuidae: Pan- 
theinae). (With J. G. Franclemont jr. author.) Proc. Entomol. Soc. Wash. 83:797-798. 

29. 1982. A new species of Zale Hiibner from Texas and New Mexico (Lepidoptera: 
Noctuidae: Catocalinae). (With J. G. Franclemont jr. author.) Proc. Entomol. Soc. Wash. 
84:134-137. 

30. 1982. Two new species of the tribe Eucosmini (Tortricidae). (With E. C. Knudson 
jr. author.) J. Lepid. Soc. 35:169-172. 

31. 1982. Two new species of Eucosma Hibner (Tortricidae) from Texas. (With E. 
C. Knudson jr. author.) J. Lepid. Soc. 35:173-178. 

32. 1982. A new species of Ozamia Ragonot (Pyralidae) from Texas. (With E. C. 
Knudson jr. author.) J. Lepid. Soc. 35:233-235. 

33. 1982. Marilopteryx carancahua, a new genus and new species from East Texas 
(Lepidoptera: Noctuidae: Hadeninae). (With J. G. Franclemont jr. author.) Proc. Entomol. 
Soc. Wash. 84:270-276. 

34. 1982. A new species of Symmetrischeria Povolny (Lepidoptera: Gelechiidae) from 
Texas. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 84:628-631. 

35. 1983. Two new species of Pyralidae (Lepidoptera) from Texas. (With E. C. Knud- 
son jr. author.) Proc. Entomol. Soc. Wash. 85:59-68. 

36. 1983. A new species of Dioryctria Zeller (Lepidoptera: Pyralidae) from Texas. 
(With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 85:116—120. 

37. 19838. A new species of Gloanna Nye (Lepidoptera: Noctuidae) from West Texas. 
(With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 85:174-176. 

38. 1983. A new genus and species of Geometridae (Lepidoptera) from Big Bend 
National Park, Texas. (With D. C. Ferguson sr. and E. C. Knudson jr. authors.) Proc. 
Entomol. Soc. Wash. 85:552-556. 

39. 1983. A new species of Psorosina Dyar (Lepidoptera: Pyralidae) from Texas. (With 
E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 85:619-621. 

40. 1983. Two new species of the tribe Eucosmini (Tortricidae) closely related to 
Phaneta granulatana (Kearfott). (With E. C. Knudson jr. author.) J. Lepid. Soc. 37:140- 
145. 

41. 1983. New North American species of Eucosmini (Lepidoptera: Tortricidae). (With 
E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 85:845-852. 


226 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


42. 1984. A new species of Hypomecis Hiibner (Lepidoptera: Geometridae) from 
Texas and Florida. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 86:291- 
294. 

43. 1984. A new Stibadium from Texas and a redescription of Striodes edentatus 
(Grote) (Noctuidae: Lepidoptera). (With E. C. Knudson jr. author.) Proc. Entomol. Soc. 
Wash. 82:346-348. 

44. 1984. Three new tortricids (Lepidoptera) from Texas. (With E. C. Knudson jr. 

-author.) Proc. Entomol. Soc. Wash. 86:446-451. 

45. 1984. A new species of Tripudia Grote (Lepidoptera: Noctuidae) from western 
Texas. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 86:639-642. 

46. 1984. A new species of Neodavisia Barnes & McDunnough (Lepidoptera: Pyral- 
idae) from southern Texas. (With D. C. Ferguson sr. and E. C. Knudson jr. authors.) Proc. 
Entomol. Soc. Wash. 86:769-772. 

47. 1984. A revision of the genus Aleptina (Lepidoptera: Noctuidae). (With E. L. 
Todd sr. and R. W. Poole jr. authors.) Proc. Entomol. Soc. Wash. 86:951-960. 

48. 1985. New species of Phycitinae (Lepidoptera: Pyralidae) from Texas, with de- 
scription of a new genus. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 87: 
231-238. 

49. 1985. A new species of Bucculatrix Zeller (Lepidoptera: Lyonetiidae) from Texas. 
(With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 87:371-374. 

50. 1985. Psychonoctua masoni (Schaus), new combination (Lepidoptera: Cossidae: 
Zeuzerinae), redescription and first records from Texas and USA. (With E. C. Knudson 
jr. author.) Proc. Entomol. Soc. Wash. 87:426—431. 

51. 1985. Two new Phycitinae (Lepidoptera: Pyralidae) from Texas and Alabama. 
(With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 87:475—479. 

52. 1985. The Eupithecia (Lepidoptera: Geometridae) of Texas, with the description 
of a new species. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 87:662— 
674. 

53. 1985. Epiblema luctuosana A. Blanchard, a homonym, is changed to Epiblema 
luctuosissima, new name. J. Lepid. Soc. 38:245. 

54. 1985. Ethmia angustalatella Powell (Lepidoptera, Oecophoridae): Description of 
the female and first U.S. records. Proc. Entomol. Soc. Wash. 87:680-681. 

55. 1985. Checklist of Lepidoptera of the Rob and Bessie Welder Wildlife Refuge 
near Sinton, Texas. (Blanchard et al.) Southwest. Entomol. 10:195-214. 

56. 1985. New U.S. records and other interesting moths from Texas. (With E. C. 
Knudson jr. author.) J. Lepid. Soc. 39:1-8. 

57. 1985. Two new species of Hexorthodes (Lepidoptera: Noctuidae) from Texas and 
Arizona. (With E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 87:777-782. 

58. 1986. Four new moths from Texas (Lepidoptera, Geometridae, Noctuidae). (With 
E. C. Knudson jr. author.) Proc. Entomol. Soc. Wash. 88:134-141. 

59. 1986. A new Chlamydastis (Oecophoridae, Lepidoptera) from Texas. (With E. C. 
Knudson jr. author.) Proc. Entomol. Soc. Wash. 88:185-188. 


NEW GENERA AND SPECIES 
Items are arranged alphabetically and keyed to the bibliography. 


Genera 


Atopothoures [=Goya Ragonot 1888] Pyralidae: Peoriinae 15, 23 
Carectocultus Pyralidae: Schoenobiinae 16 

Glyphocystis Pyralidae: Phycitinae 13 

Pimodes_ Pyralidae: Phycitinae 21 

Pseudocabotia Pyralidae: Phycitinae 48 

Rostrolaetilia Pyralidae: Phycitinae 17 

Triozosneura Pyralidae: Phycitinae 13 

Welderella Pyralidae: Phycitinae 24 


VOLUME 41, NUMBER 4 : 227 


Species 


adusta, Euamiana Noctuidae 58 
TL: Texas, Jeff Davis Co., Davis Mountains, Madera Canyon. Holotype (HT) 4, 17 
August 1984, in USNM. 
albisericea [albisericeum], Copablepharon Noctuidae 19 
TL: Texas, Hemphill Co., nr. Canadian, Gene Howe Wildlife Management Area. HT 
6, 27 September 1968, in USNM (type no. 73431). 
anaimella, Meroptera Pyralidae 48 
TL: Texas, Presidio Co., Shafter. HT 6, 9 July 1969, in USNM. 
apicigrammella, Melitara Pyralidae 48 
TL: Texas, Terrell Co., Sanderson. HT 6, 28 September 1980, in USNM. 
arenella, Eoreuma_ Pyralidae 35 
TL: Texas, [Kleberg Co.], Padre Island National Seashore. HT 4, 19 July 1976, in USNM. 
argutipunctana, Phaneta Tortricidae 40 
TL: Texas, Hemphill Co., Canadian. HT 6, 15 August 1971, in USNM. 
atascosana, Eucosma_ Tortricidae 26 
TL: Texas, Cameron Co., Laguna Atascosa National Wildlife Refuge. HT 6, 22 No- 
vember 1973, in USNM (type no. 75821). 
atratella, Salebriaria Pyralidae 51 
TL: Texas, Hunt Co., Lake Twakoni, Wind Point Park. HT 4, 15 July 1984, in USNM. 
auripurpura, Hydroecia Noctuidae 4 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch, 1613 m. HT 6, 11 
October 1966, in USNM (type no. 68160). 
balconiensis, Pseudocabotia Pyralidae 48 
TL: Texas, Kerr Co., 10 miles west of Hunt. HT 6, 4 September 1981, in USNM. 
benitensis, Heterocampa Notodontidae 7 
TL: Texas, Cameron Co., Brownsville. HT 6, 8 August 1967, in USNM (type no. 64647). 
bucurvata, Astalotesia Geometridae 38 
TL: Texas, Brewster Co., Big Bend National Park, Chisos Basin. HT 6, 29 March 1982, 
in USNM. 
caelebs, Pyromorpha Zygaenidae 10 
TL: Texas, Jeff Davis Co., Fort Davis, Hospital Canyon. HT 6, 18 May 1971, in USNM 
(type no. 71981). 
caesirufella, Dioryctria Pyralidae 36 
TL: Texas, Kerr Co., Kerrville State Park. HT 4, 19 August 1980, in USNM. 
caesium, Stibadium Noctuidae 43 
TL: Texas, Cameron Co., south Padre Island. HT 4, 24 October 1982, in USNM. 
callaisata, Paramiana Noctuidae 9 
TL: [Texas, Culberson Co.], Guadalupe Mountains [National Park], Pine Spring Canyon, 
1737 m. HT 4, 28 August 1967, in USNM (type no. 68149). 
carancahua, Marilopteryx Noctuidae 33 
TL: Texas, Jackson Co., Deutschburg nr. Carancahua Creek. HT 6, 6 March 1975, in 
USNM. 
chihuahua, Tripudia Noctuidae 45 
TL: Texas, Brewster Co., Big Bend National Park, Chihuahuan Desert nr. Nugent 
Mountain. HT 6, 8 October 1969, in USNM. 
chisosensis, Zale Noctuidae 29 
TL: Texas, Jeff Davis Co., Davis Mountains, Mt. Locke, 2042 m. HT 6, 10 June 1969, 
in USNM. 
citeria, Hexorthodes Noctuidae 57 
TL: Texas, Jeff Davis Co., Fort Davis. HT 6, 11 June 1969, in USNM. 
clarkei, Phaneta Tortricidae 41 
TL: Texas, Hemphill Co., Canadian National Grassland, Lake Marvin. HT ¢, 9 October 
1982, in USNM. 
collilonga, Pelochrista Tortricidae 44 
TL: Texas, Brown Co., Lake Brownwood State Park. HT 6, 21 April 1966, in USNM. 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


coloradella, Rostrolaetilia Pyralidae 17 
TL: Colorado, [Pueblo Co.], Pueblo. HT 2, July, in USNM (type no. 73284). 
cottami, Oncocnemis Noctuidae 9 
TL: Texas, [Brewster Co.], Big Bend National Park, Basin, 1676 m. HT 6, 10 May 1966, 
in USNM. 
cruentana, Phaneta Tortricidae 30 
TL: Texas, Anderson Co., nr. Tennessee Colony, [Gus] Engeling Wildlife Management 
Area. HT 6, 28 June 1978, in USNM (type no. 76733). 
diabolana, Eucosma_ Tortricidae 26 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area, 1829 m. HT 8, 
31 March 1970, in USNM (type no. 75820). 
dominicki, Carectocultus Pyralidae 16 
TL: Texas, Jackson Co., Deutschburg. HT 4, 31 July 1972, in USNM (type no. 73242). 
dorsonotata, Triozosneura Pyralidae 13 
TL: Texas, [Jeff Davis Co.], Davis Mountains, Mt. Locke, McDonald Observatory grounds. 
HT 6, 27 August 1970, in USNM (type no. 72379). 
dupla, Bertelia Pyralidae 20 
TL: Texas, Presidio Co., Shafter. HT 4, 19 October 1973, in USNM (type no. 73530). 
emendata, Hexorthodes Noctuidae 57 
TL: Texas, Jeff Davis Co., Fort Davis. HT 6, 11 June 1969, in USNM. 
eureka, Rostrolaetila Pyralidae 17 
TL: Utah, Eureka. HT 4, 14 August 1911, in USNM (type no. 73285). 
exculta, Paramiana Noctuidae 58 
TL: Texas, Jeff Davis Co., Mount Locke. HT ¢, 20 August 1984, in USNM. 
fergusonella, Psorosina Pyralidae 39 
TL: Texas, Anderson Co., nr. Tennessee Colony, [Gus] Engeling Wildlife Management 
Area. HT 6, 19 June 1982, in USNM. 
franclemonti, Oxycnemis Noctuidae 4 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch, 1646 m. HT 4, 3 
April 1965, in USNM (type no. 68162). 
fritillana, Eucosma_ Tortricidae 30 
TL: Texas, Anderson Co., nr. Tennessee Colony, [Gus] Engeling Wildlife Management 
Area. HT 4, 28 June 1978, in USNM (type no. 76734). 
furtiva, Ursia Notodontidae 8 
TL: [Texas, Brewster Co.], Big Bend National Park, Pine Canyon, 1585 m. HT 4, 2 
September 1964, in USNM (type no. 64648). 
garneri, Drepanulatrix Geometridae 58 
TL: Texas, Uvalde Co., Garner State Park. HT 6, 24 March 1985, in USNM. 
gillaspyi, Copablepharon Noctuidae 19 
TL: Texas, Kleberg Co., Padre Island National Seashore. HT 6, 28 September 1978, in 
USNM (type no. 734383). 
graziella, Eucosma Tortricidae 4, 26 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch. HT 6, 11 October 
1966, in USNM (type no. 68164). 
griselda, Eucosma Tortricidae 31 
TL: Texas, Brewster Co., Big Bend National Park, Chisos Basin. HT 6, 7 April 1967, 
in USNM. 
guttulana, Eucosma_ ‘Tortricidae 26 
TL: Texas, Kleberg Co., Padre Island National Seashore. HT 4, 19 July 1976, in USNM 
(type no. 75819). 
habrolepis, Chlamydastis Oecophoridae 59 
TL: Texas, Cameron Co., Laguna Atascosa [National Wildlife Refuge]. HT 6, 1 April 
1978, in USNM. 
hecate, Gloanna Noctuidae 37 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area. HT 4, 11 June 
1982, in USNM. 


VOLUME 41, NUMBER 4 229 


heppneri, Petrophila Pyralidae 35 
TL: Texas, Kerr Co., [16.1 km] W of Hunt. HT 6, 1 September 1980, in USNM. 
heterogena, Oncocnemis Noctuidae 9 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch. HT 6, 27 August 
1965, in USNM (type no. 68148). 
hieroglyphana, Grapholita Tortricidae 44 
TL: Texas, [Culberson Co.], Guadalupe Mountains, Nickel Creek. HT 4, 10 July 1968, 
in USNM. 
insularis, Pimodes_ Pyralidae 21 
TL: Texas, Kleberg Co., Padre Island National Seashore. HT 6, 29 September 1975, in 
USNM (type no. 73652). 
junctimacula, Aleptina Noctuidae 47 
TL: Texas, [Brewster Co.], Big Bend National Park, Dugout Wells. HT 6, 29 August 
1965, in USNM. 
kendalli, Bucculatrix Lyonetiidae 49 
TL: Texas, Bexar Co., Ebony Hill Research Station (Kendall residence). HT 4, 19 
September 1984, in USNM. 
kendalli, Zamagiria Pyralidae 6 
TL: Texas, Jeff Davis Co., Fort Davis, Hospital Canyon, 1524 m. HT 4, 11 July 1969, 
in USNM (type no. 71004). 
kendallorum, Symmetrischema_ Gelechiidae 34 
TL: Texas, Nueces Co., north Padre Island. HT 6, 17 September 1981, in USNM. 
lenticuligera, Glenoides Geometridae 12 
TL: Texas, Hidalgo Co., Santa Ana National Wildlife Refuge. HT 6, 15 February 1971, 
in USNM (type no. 72326). 
linitipunctana, Phaneta Tortricidae 40 
TL: Texas, Nueces Co., north Padre Island. HT 6, 9 September 1974, in USNM. 
longipectinaria, Hypomecis Geometridae 42 
TL: Texas, Montgomery Co., Conroe. HT 6, 30 April 1970, in USNM. 
luctuosana, Epiblema Tortricidae (homonym—see luctuosissima) 25, 53 
luctuosissima, Epiblema Tortricidae 25, 53 
TL: Texas, Nueces Co., north Padre Island. HT 6, 6 April 1978, in USNM (type no. 
75822). 
margueritaria, Grotella Noctuidae 4 
TL: Texas, [Brewster Co.], Big Bend National Park, Chihuahuan Desert nr. Nugent 
Mountain, 914 m.HT 4, 8 October 1966, in USNM (type no. 68168). 
mayelisana, Phaneta Tortricidae 26 
TL: Texas, Cottle Co., nr. Paducah, Matador Wildlife Management Area. HT 4, 17 
April 1968, in USNM (type no. 75817). 
mayelisaria, Glaucina Geometridae 3 
TL: [Texas, Brewster Co.], Big Bend National Park, Government Spring. HT 4, 29 
September 1965, in AMNH. 
melusina, Neodavisia Pyralidae 46 
TL: Texas, Starr Co., Roma. HT 6, 4 April 1978, in USNM. 
mendaciana, Suleima Tortricidae 41 
TL: Texas, Brewster Co., Big Bend National Park, Dugout Wells. HT 6, 28 September 
1981, in USNM. 
mephisto, Neperigea [Properigea| Noctuidae 4 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area, Sierra Diablo 
Mountains, NNW of Van Horn, 1676 m. HT 4, 22-23 June 1965, in USNM (type no. 
68161). 
minimella, Rostrolaetilia Pyralidae 17 
TL: California, Inyo Co., Olancha. HT 4, “June 24-30” in USNM (type no. 73281). 
multistriatella, Ozamia_ Pyralidae 32 
TL: Texas, Jeff Davis Co., Fort Davis. HT 6, 25 March 1968, in USNM. 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


musetta, Phaneta Tortricidae 41 
TL: New Mexico, Socorro Co., Gran Quivara National Monument, 2012 m. HT 4, 1- 
3 July 1964, in USNM. 
nymphana, Gretchena_ Tortricidae 41 
TL: Texas, Anderson Co., nr. Tennessee Colony, Gus Engeling Wildlife Management 
Area. HT 4, 15 April 1968, in USNM. 
_ ovaliger, Atopothoures [=Goya Ragonot 1888] Pyralidae 15, 23 
TL: Texas, [Kimble Co.], Junction. HT 6, 21 August 1978, in USNM (type no. 73241). 
padreella, Peoria FPyralidae 27 
TL: Texas, Kleberg Co., Padre Island National Seashore. HT 4, 24 June 1976, in USNM 
(type no. 76140). 
paraplesiana, Sonia Tortricidae 25 
TL: Texas, [Harris Co.], Houston. HT 6, 5 June 1968, in USNM. 
parvalbum, Homoeosoma Pyralidae 48 
TL: Texas, Brewster Co., Big Bend National Park, Hot Springs. HT 6, 4 April 1984, in 
USNM. 
peorinella, Anderida Pyralidae 51 
TL: Texas, Brewster Co., Big Bend National Park, K-Bar Research Station. HT 4, 1 
April 1984, in USNM. 
pinalensis, Rostrolaetilia Pyralidae 17 
TL: Arizona, [Gila Co.], Pinal Mountains, 1524 m. HT 4, 15-30 April 1925, in USNM 
(type no. 73287). 
placidissima, Rostrolaetilia Pyralidae 17 
TL: Utah, Stockton. HT 6, “IX.1.4”, in USNM (type no. 73282). 
salaciana, Eucosma_ Tortricidae 31 
TL: Texas, Nueces Co., north Padre Island. HT 6, 13 October 1979, in USNM. 
salmocolor, Dasypyga Pyralidae 6 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area, 1829 m. HT 4, 1 
September 1969, in USNM (type no. 71005). 
septuosa, Tarachidia Noctuidae 58 
TL: Texas, Cameron Co., Laguna Atascosa [National Wildlife Refuge]. HT 4, 16 May 
1974, in USNM. 
serraticornis [serraticorne], Copablepharon Noctuidae 19 
TL: Texas, Cottle Co., nr. Paducah, Matador Wildlife Management Area. HT 4, 8 
September 1966, in USNM (type no. 734382). 
sierrae, Eucosma_ Tortricidae 41 
TL: Texas, Culberson Co., Sierra Diablo Wildlife Management Area, 1951 m. HT 4, 30 
August 1970, in USNM. 
signifera, Macrorrhinia Pyralidae 21 
TL: Texas, Tyler Co., Town Bluff. HT 6, 7 August 1975, in USNM (type no. 73651). 
texanalis, Oenobotys Pyralidae 22 
TL: Texas, Jeff Davis Co., Fort Davis. HT 6, 5 October 1969, in CNC (type no. 13919); 
allotype-topotype ¢ in USNM. 
texanella, Rostrolaetilia Pyralidae 17 
TL: Texas, Jeff Davis Co., Davis Mountains, Mt. Locke. HT 4, 4 July 1969, in USNM 
(type no. 73286). 
texasana, Anopina Tortricidae 44 
TL: Texas, Jeff Davis Co., [Davis Mountains], Mt. Locke, 2042 m. HT 4, 26 April 1981, 
in USNM. 
toddi, Oncocnemis Noctuidae 4 
TL: Texas, [Brewster Co.], Big Bend National Park, Chihuahuan Desert nr. Dugout 
Wells, 914 m. HT 6, 28 September 1965, in USNM (type no. 68165). 
ustulatana, Eucosma Tortricidae 41 
TL: Texas, Washington Co., Brenham. HT 4, 4 June 1979, in USNM. 
utahensis, Rostrolaetilia Pyralidae 17 
TL: Utah, Richfield. HT ¢, 15 June 1930, in USNM (type no. 73288). 


VOLUME 41, NUMBER 4 231 


valliscola, Acronicta Noctuidae 4 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch, 1585 m. HT 4, 10 
May 1966, in USNM (type no. 68159). 

verecundana, Phaneta Tortricidae 26 
TL: Texas, Hemphill Co., Gene Howe Wildlife Management Area nr. Canadian. HT 
6, 15 August 1971, in USNM (type no. 75818). 

viridivallis, Glyphocystis Pyralidae 13 
TL: Texas, [Brewster Co.], Big Bend National Park, Green Gulch. HT é, 28 March 
1971, in USNM (type no. 72380). 


RARE AND INTERESTING LEPIDOPTERA 
Species are arranged alphabetically and keyed to the bibliography. 


addens, Eulepidotis Noctuidae (first TX & US, ? fig.) 56 

admixta, Quasisalebria Pyralidae (first TX) 6 

alatella, Myelopsis Pyralidae (first TX) 6 

albiplagiatella occidentalis, Pima Pyralidae (first TX) 6 

aleptivoides, Aleptina Noctuidae (new comb., 6, 6 & 2 genit. fig.) 47 

alpinata, Eupithecia Geometridae (6 & genit. fig.) 52 

anartoides, Radara {[Matiloxis] Noctuidae (first TX & US (?), 6 fig.) 11 

angustalatella, Ethmia Oeccophoridae (first TX & US, 6, 2 & genit. fig.) 54 

ardiferella, Rostrolaetilia Pyralidae (HT 2, 6 & 2 genit. fig.) 17 

arizonensis, Daulia Pyralidae (6 fig.) 56 

atalanta, Scordylia [Heterosia] Geometridae (first TX & US (?), 2 fig.) 11 

auranticella(=rossi, corrected in #14), Dioryctria Pyralidae (first TX) 6 

australelia, Zamagiria Pyralidae (TX TL conf.) 6 

baracoalis, Lamprosema_ Pyralidae (2 fig.) 56 

basipunctaria, Sigela Noctuidae (first TX (?), 6 fig.) 11 

belfragei, Heterocampa Notodontidae (comp. with H. benitensis) 7 

bolteri, Eupithecia Geometridae (6 & genit. fig.) 52 

brillians, Miracavira Noctuidae (6 fig.) 56 

cana, Sparganothis Tortricidae (6 fig.) 56 

clinopetes, Aleptina Noctuidae (new comb., 6, 6 & 2 genit. fig.) 47 

cocoata, Eupithecia Geometridae (first TX) 52 

coloradensis, Eupithecia Geometridae (first TX) 52 

collaris, Nystalea Notodontidae (larval foodplant, 2 fig.) 56 

concors, Glympis Noctuidae (TX records, ¢ fig.) 11 

condigna, Anadelosemia_ Pyralidae (first TX) 6 

consimilis, Evergestis Pyralidae (TX records, é fig.) 11 

costata, Stemhorrhages Pyralidae (6 fig.) 56 

cucullidea, Emariannia Noctuidae (TX records, 6 fig.) 11 

daemonalis, Iscadia Noctuidae (TX records, é fig.) 11 

dissimulatrix, Actrix Pyralidae (first TX) 6 

donysa, Amplypterus Sphingidae (first TX & US, @ fig.) [=Adhermarius blanchar- 
dorum Hodges 1985] 11 

eburneata, Scopula Geometridae (6 fig.) 56 

edentatus, Stiriodes Noctuidae (first TX, 6 fig.) 43 

edna, Eupithecia Geometridae (6 & genit.) 52 

electrica, Goniocarsia Noctuidae (@ fig.) 56 

erronella, Unadilla Pyralidae (first TX & US) 6 

eudoreella, Diviana Pyralidae (6 fig.) 56 

formularis, Lesmone Noctuidae 56 

fredericki, Eupithecia Geometridae (OD, 6 & genit. fig.—Knudson) 52 

fufius, Lesmone Noctuidae (6 fig.) 56 

funeralis, Acronicta Noctuidae (first TX, 6 fig.) 56 

grandis [grande], Copablepharon Noctuidae (TX records) 19 


232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


granitella, Pima Pyralidae (first TX) 6 

graziella, Eucosma_ Tortricidae (2 genit. fig.) 26 

grisella, Bertelia Pyralidae (first TX) 6 

heiligbrodti, Syssphinx [Sphingicampa] Saturniidae (larval foodplants) 2 
huachuca, Eupithecia Geometridae (6 & genit. fig.) 52 

idella, Hemispraguei Noctuidae (6 fig.) 56 

_ ignobilis, Fundella Pyralidae (first TX & US) 6 

impressale, Homoeosoma Pyralidae (first TX) 6 

inca, Aleptina Noctuidae (generic rev., 6 & genit. fig.) 47 

ingenua, Charadra Noctuidae (first TX, 6, 6 & 2 genit.) 28 

inornata, Episcepsis Ctenuchidae (first TX & US (?), 2 fig.) 11 

insignis, Eremberga Pyralidae (first TX & US) 6 

iole, Eriopyga Noctuidae (first TX & US, é fig.) 56 

jejunata, Eupithecia Geometridae (6 & genit. fig.) 52 

kearfottella, Acrobasis Pyralidae (first TX) 6 

libya, Xylophanes Sphingidae (first TX & US, 6 fig.) 56 

longidens kerrvillaria, Eupithecia Geometridae (6 genit. fig.) 52 

maestosa, Eupithecia Geometridae (first TX) 52 

masoni, Psychonoctua_ Cossidae (new comb., first TX & US, 6 & genit. fig.) 50 
matheri, Eupithecia Geometridae (6 & genit. fig.) 52 

maximellus, Schoenobius Pyralidae (life hist., larval foodplant, larva & pupa fig.) 1 
melanthus, Syntomeida Ctenuchidae (first TX & US (?), 6 fig.) 11 
mexicanaria, Phaeoura Geometridae (gynandromorph, adult & genit. fig.) 5 
minutana, Epiblema_ Tortricidae [syn. of strenuana] (rev. status) 25 
miserulata, Eupithecia Geometridae (6 & genit. fig.) 52 

mulina, Eriopyga [Protorthodes] Noctuidae (first TX & US (?), 9 fig.) 11 
nigromaculella, Rostrolaetilia Pyralidae (HT 9, 6 & 2 genit. fig.) 17 
nonparilella, Sosipatra Pyralidae (2 fig.) 56 

nyssaecolella, Actrix Pyralidae (first TX) 5 

obtusus [obtusa], Stiriodes Noctuidae (comp. with S. edentatus) 43 

olivata, Aeschropteryx [Prochoerodes| Geometridae (first TX & US (?), 6 fig.) 11 
ornata, Eupithecia Geometridae (6 & genit. fig.) 52 

packardella, Rhagea Pyralidae (first TX) 6 

palmata, Hypopta Cossidae (6 fig.) 56 

parallela, Hemiplatytes Pyralidae (6 fig.) 56 

parvella, Ollia Pyralidae (to new genus Welderella) 24 

peckorum, Eupithecia Geometridae (6 & genit. fig.) 52 

persimulata, Eupithecia Geometridae (6 & genit. fig.) 52 

pertusata, Eupithecia Geometridae (6 & genit. fig.) 52 

pharaxalis, Bocchoropsis Pyralidae (first TX & US, 2 fig.) 56 

pimensis, Euxoa Noctuidae (first TX, 6 fig.) 56 

placidata, Eupithecia Geometridae (6 & genit. fig.) 52 

placidella, Rostrolaetilia Pyralidae (LT 4, 6 & 2 genit. fig.) 17 

prolongalis, Microthyris Pyralidae (6 fig.) 56 

prona, Orthogramma [Epitausa] Noctuidae (first TX & US (?), 6 fig.) 11 
prostrata, Eupithecia Geometridae (6 & genit. fig.) 52 

psegmapteryx, Matigramma[Toxonprucha] Noctuidae (first TX & US(?), éfig.) 11 
pusillata interruptofasciata, Eupithecia Geometridae (TX from type only) 52 
redtenbacheri, Givira [Comadia] Cossidae (first TX & US, 4 fig.) 11 
respondens, Neophanis Noctuidae (first TX & US (?), 2 fig.) 11 

ridingsana, Eucosma_ Tortricidae (comp. with E. griselda) 29 

rosea, Oncocnemis Noctuidae (first TX, 6 fig.) 56 

rossi, Dioryctria Pyralidae (first TX) 14 

ruthaea, Cropia Noctuidae (first TX & US, 6 fig.) 56 

sanguinalis, Polygrammodes_ Pyralidae (first TX & US, 6 fig.) 11 

semiatra, Aleptina Noctuidae (new comb., 6, 6 & 2 genit. fig.) 47 

semicana, Patriciola’ Pyralidae (first TX) 6 


VOLUME 41, NUMBER 4 233 


serrata [serratum], Copablepharon Noctuidae (comp. with C. serraticornis & C. gil- 
laspyi) 19 

servia, Rhescipha [Goniapteryx] Noctuidae (first TX (?), ¢ fig.) 11 

sicheas, Gonodonta Noctuidae (first TX & US(?)) 11 

sierrae, Eupithecia Geometridae (6 & genit. fig.) 52 

sinaldus, Gonodonta Noctuidae (first TX & US (?), 6 fig.) 11 

sincera, Salobrena_ Pyralidae (6 fig.) 56 

sonorella, Anderida Pyralidae (first TX) 6 

stercorea, Ancylostomia Pyralidae (first TX) 6 

stigmaphiles, Herminodes [Rhosologia] Noctuidae (first TX & US (?), 2 fig.) 11 

stigmella, Acrobasis  Pyralidae (first TX) 6 

subulalis, Araschnopsis Pyralidae (first TX & US, 9? fig.) 56 

superba, Heterocampa Notodontidae (comp. with H. benitensis) 7 

swettii, Eupithecia Geometridae (6 & genit. fig.) 52 

terminalis, Oncocnemis Noctuidae (first TX, 6 fig.) 56 

titan, Meropleon Noctuidae (first TX & US (?), 6 fig.) 11 

valta, Pentobesa Notodontidae (first TX & US (?), 6 fig.) 11 

wellingtoniana, Anopina Tortricidae (LT desig.) 44 

woodgatata, Eupithecia Geometridae (é & genit. fig.) 52 

xasta, Euxoa Noctuidae (first TX (?), 6 fig.) 11 

xylia, Letis Noctuidae (first TX & US, 6 fig.) 56 

xygadeniata, Eupithecia Geometridae (6 & genit. fig.) 52 


INTENSIVELY WORKED COLLECTING LOCALITIES 


Although some collections were made in other states (Colorado and New Mexico), Texas 
was of primary interest and intensively worked by Blanchard. In early 1962 he began 
preparing a card index of species found in Texas. Taxonomic arrangement was primarily 
that of McDunnough’s 1938 and 1939 Check Lists. For each of more than 3000 species 
he cited the original description, other significant taxonomic references, and listed specific 
records including his own captures. The card file was designed for his personal use and 
as a memory jogger. Locality data rarely included details. 

Because these localities apply to approximately 100,000 specimens in various museums 
and private collections, and because future workers will be interested in specific localities 
for spatial distribution and life history studies, more complete locality data are provided 
here. During one of our last visits André permitted me to make photocopies of his 
“Lepidoptera Card File” for my own data-bank. It was his wish, however, that the file 
eventually go to Edward C. Knudson, coauthor of about half of his published papers. It 
did. Collections were made in more than 57 counties; there are 254 in the State, thus the 
sample size was 22% or more of the total. 

Locations are arranged alphabetically, and each includes Texas county and other details. 


Addicks. Harris Co., W of Houston off [H-10. 

Aguja Canyon. Jeff Davis Co., Davis Mountains, Tx Ranch road 1832, 18 km W of Tx 
Hwy 17 at Boy Scout Camp. 

Alpine. Brewster Co., within city of Alpine or nearby. 

Alpine Girl Scout Camp. Jeff Davis Co., off Tx Hwy 118, 23 km NNW of Alpine nr. 
Miter Peak. 

Aransas National Wildlife Refuge. Aransas Co., 11 km S of Austwell (133,982+ ha). 
Artesia Wells. Dimmit-LaSalle counties, Chaparral Wildlife Management Area, N of 
Farm Road 183, 8 km W of Artesia Wells. (Collections mostly in LaSalle Co.) 

Balmorhea State Park. Reeves Co., Tx Hwy 17 nr. Toyahavale. 
Bastrop State Park. Bastrop Co., Tx Hwy 21 nr. Bastrop. 
Bear Canyon. (See Guadalupe Mountains National Park.) 
Bear Creek. Harris Co., Bear Creek Park, recreation area in NW Houston. 
Belton Reservoir. Bell Co., Lake Belton nr. Belton. 
Big Bend National Park. Brewster Co. (1,746,997 ha) 
Basin 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Chihuahuan Desert nr. Nugent Mountain (Nugent Mt. draw) 
Chisos Mountains (principal mts. within Park) 
Croton Spring 
Dugout Wells 
Government Spring 
Grapevine Hills & Spring 
Green Gulch 
K-Bar Research Station (old ranch house nr. Hq.) 
Oak Spring 
Old Ranch (Burro Messa, road to Castilon) 
Panther Canyon (draw, spring) below Park Hq. 
Panther Junction 
Pine Canyon 
Rio Grande Village 
San Vicente Crossing 
(All locations indicated on maps available at Park Hq.) 
Black Gap. Brewster Co., Black Gap Wildlife Management Area, Ranch road 2627, 56 
km SE of Marathon (254,513+ ha). 
Blanco State Park (or only Blanco). Blanco Co., on Blanco River at Blanco city. 
Brazos River. (See Spivey Crossing.) 
Caddo Lake State Park (or only Caddo Lake). Harrison Co., nr. Kanak. 
Camp Strake. (See Conroe.) 
Canadian. (See Gene Howe Wildlife Management Area.) 
Chihuahuan Desert nr. Nugent Mt. (See Big Bend National Park.) 
Churchill Bridge. Brazoria Co., SW of Brazoria off Tx Hwy 36 at San Bernard River. 
Clebourne State Park (or only Cleburne). Johnson Co., SW of Clebourne off US Hwy 67. 
Coldspring. San Jacinto Co., Sam Houston National Forest, Tx Hwy 2025, 5 km S of 
Coldspring, Double Lake recreation area. 
College Station. Brazos Co., joins city of Bryan. 
Conroe. Montgomery Co., Camp Strake, private recreation area at Conroe. 
Cypress & Cypress Lake. Harris Co., NW of Houston, US Hwy 290/Tx Hwy 6. 
Dam B (now Steinhagen Lake). Tyler Co., 29 km E of Woodville on Neches River. 
(Traps were usually set near the spillway or along W bank of reservoir.) 
Deussen Park. Harris Co., Alexander Deussen Park, Houston (N Lake Houston Parkway 
on Lake Houston). 
Deutschburg. Jackson Co., a community nr. Carancahua Creek S of Francitas. 
Dewalt. Fort Bend Co., small community on Tx Hwy 6. 
Don George Lake. Fort Bend Co., 16 km SE of Richmond. 
Eagle Lake. Colorado Co., US Hwy 90-A. 
Eagle Pass. Maverick Co., private property nr. Eagle Pass. 
El Rancho Cima. Comal/Hays counties, Tx Hwy 32, 24 km W of San Marcos, Boy 
Scout Camp. (Most collections were made in Hays Co.) 
Falcon Heights. Zapata Co., nr. Falcon Reservoir on Rio Grande. 
Falcon State Park. Zapata Co., off US Hwy 83 on Falcon Reservoir. 
Frijoles. (See Guadalupe Mountains National Park.) 
Garner State Park (or only Garner). Uvalde Co., off US Hwy 83 40 km N of Uvalde. 
Gene Howe Wildlife Management Area. Hemphill Co., FM road 2266, 10 km E of 
Canadian (14,383+ ha). 
George West. Live Oak Co., private property nr. George West. 
Goodrich. Polk Co., S of Livingston. (Collection in Sam Houston National Forest.) 
Gray Ranch. (See Jeff Davis Co.) 
Guadalupe Mountains National Park. Culberson & Hudspeth counties off US Hwy 62/ 
180 (191,546+ ha). (All collections made in Culberson Co.) 
Bear Canyon 
Frijoles, nr. Manzanita Spring 
McKittrick Canyon 


VOLUME 41, NUMBER 4 235 


Pine Spring Canyon (or only Pine Canyon) 
Smith Canyon 
(Local National Park map shows all locations.) 
Guadalupe River. Comal Co., on Guadalupe River NNW of New Braunfels, private 
recreation camp nr. 4th crossing of river. 
Gus Engeling Wildlife Management Area. Anderson Co., US Hwy 287, 32 km NW of 
Palestine nr. Tennessee Colony (26,392+ ha). 
Harlingen. Cameron Co., private property nr. Harlingen. 
Hempstead. Waller Co., US Hwy 290/Tx Hwy 6. 
Huntsville State Park (or only Huntsville). Walker Co., off Tx Hwy 19, 8 km S of 
Huntsville. 
Houston. Harris Co., unless otherwise indicated, 3025 Underwood Street, Blanchard’s 
residence. 
Indian Village. Polk Co., Alabama & Coushatta Indian Reservation, off US Hwy 190, 
24 km E of Livingston. 
Jeff Davis Co. 
Fort Davis 
Hospital Canyon, behind Old Fort Davis Historic Site 
Limpia Canyon, Tx Hwys 17 & 118 nr. Fort Davis 
Mt. Livermore, off Tx Hwy 166, 8 & 13 km SE of, on Friend’s Ranch (private) 
Tom R. Gray Ranch, Tx Hwy 166 WSW of Fort Davis (private) 
W. B. Sharp Ranch, Tx Hwy 17 NNE of Fort Davis (private) 
Junction. Kimble Co., V. H. Ranch, US Hwy 377, 14 km SW of Junction on South Llano 
River. 
Kerrville State Park (or only Kerrville). Kerr Co., on Guadalupe River at Kerrville. 
Kerr Wildlife Management Area. Kerr Co., Tx Ranch road 1340, 45 km W of Kerrville 
(16,044+ ha). 
Laguna Atascosa. (See Laguna Atascosa National Wildlife Refuge.) 
Laguna Atascosa National Wildlife Refuge. Cameron/Willacy counties on Laguna Madre 
(111,558+ ha). (All collections made in Cameron Co.) 
Laguna Park. Hill Co., small town, collections on grounds of local motel. 
Lake Brownwood State Park. Brown Co., off Tx Hwy 279, 26 km NNW of Brownwood. 
Lake Corpus Christi State Park. San Patricio Co., off Tx Hwy 359, 6 km SW of Mathis. 
Lake o the Pines. Marion Co., private recreation area, N side of lake. 
Lake Travis. Travis Co., recreation area on Lake Travis nr. Bee Cave off Tx Hwy 71. 
Lake Walk. Val Verde Co. Before 1969 this and Devils Lake were flood control reservoirs 
on the Devils River I] and 16 km above the Rio Grande. Following completion of 
Amistad Dam and Reservoir on the Rio Grande, both Lake Walk and Devils Lake 
ceased to exist, having been inundated by water from Amistad. 
Livingston. Polk Co. 
Matador Wildlife Management Area. Cottle Co., W of US Hwy 838, 18 km NW of 
Paducah (69,242+ ha). 
Mathis. San Patricio Co., private property nr. Mathis. 
McKittrick Canyon. (See Guadalupe Mountains National Park.) 
Memorial Park. Harris Co., Houston city park. 
Morgan's Point. Harris Co., E of Pasadena on San Jacinto Bay. 
Mount Locke. Jeff Davis Co., Davis Mountains, off Tx Hwy 118, 26 km NW of Fort 
Davis, grounds of McDonald Observatory. 
New Waverly. Walker Co., Sam Houston National Forest, Stubblefield recreation area 
W of New Waverly. 
Nickel Creek. Culberson Co., US Hwy 62/180, a small community 8 km NE of Pine 
Springs. (Collections made in Guadalupe Mountains National Park.) 
Padre Island (Tx offshore island spanning 5 counties). 
North Padre Island. Nueces Co. 
Padre Island National Seashore. Kleberg and Kenedy counties. (Collections made in 
Kleberg Co.) 


236 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


South Padre Island. Willacy Co. (north) and Cameron Co. (south). (Collections made 

in Cameron Co.) 

Paducah. (See Matador Wildlife Management Area.) 

Palo Duro Canyon State Park. Randall Co., Tx Hwy 217, 19 km E of Canyon. 

Pedernales Falls State Park. Blanco Co., Tx Farm road 2766, 13 km E of Johnson City 
on Pedernales River. 

_ Point Comfort. Jackson Co., private property on Lavaca Bay. 

Port Alto. Calhoun Co., W shore Carancahua Bay. Blanchard had a beach house here 
destroyed by Hurricane Celia 3 August 1970. 

Port Lavaca. Calhoun Co., on Lavaca Bay. 

Possum Kingdom. Palo Pinto Co., Possum Kingdom State Park, on W shore of Possum 
Kingdom Reservoir. 

Raymondville. Willacy Co., private property nr. Raymondville. 

Richmond. Fort Bend Co., private property, E bank of Brazos River off US Hwy 59. 

Rio Frio. Real Co., small village on Frio River off US Hwy 83, 13 km S of Leakey. 

Rosenberg. Fort Bend Co., nr. Rosenberg (private). 

Ruidosa-Hot Spring. Presidio Co., 10 km ENE of Ruidosa on Hot Springs Creek, private 
recreation area. 

Sabinal. Uvalde Co., private recreation area nr. Sabinal. 

San Antonio. Bexar Co., Mountain View Acres, 5598 Mt. McKinley Drive NE (R. O. & 
C. A. Kendall residence which they dubbed Ebony Hill Research Station, “EHRS’). 

San Benito. Cameron Co., private property nr. San Benito. 

Santa Ana Refuge (or Santa Ana). Hidalgo Co., Santa Ana National Wildlife Refuge, 
off US Hwy 281 S of Alamo (4885+ ha). 

Santa Rosa. Cameron Co., Las Palomas Wildlife Management Area—Longoria Tract, 
Tx Farm road 506, 6 km N of Santa Rosa. 

Sealy. Austin Co., US Hwy 90 at Brazos River (private). 

Shafter. Presidio Co., private property, recreation area. 

Sharp Ranch. (See Jeff Davis Co.) 

Sheffield. Pecos Co., Tx Hwy 349 at US Hwy 290 nr. Pecos River. 

Sheldon. Harris Co., Sheldon Wildlife Management Area, US Hwy 90, 16 km E of 
Houston (6046+ ha). 

Shepherd. San Jacinto Co., small community on US Hwy 59. (Collections in Sam Houston 
National Forest.) 

Sierra Diablo. Culberson Co., Sierra Diablo Wildlife Management Area, 51 km NW of 
Van Horn (19,251+ ha). 

Smith Point. Chambers Co., small community on E shore of Trinity Bay. 

Spivey Crossing. Hill Co., 11 km SE of Laguna Park, Tx Farm road 2114 at Brazos 
River, J. W. Glenn Ranch. 

Spring. Harris Co., small town on US Hwy 75 N of Houston (private property). 

Tennessee Colony. Collections made at Gus Engeling Wildlife Management Area. 

Town Bluff. Tyler Co., small village adjacent to Steinhagen Lake dam (formerly Dam 
B), Tx Farm road 92. 

Utopia. Uvalde Co., Tx Hwy 187, NE corner of county, 2 km S of Utopia, Old Waresville 
River Resort. 

Van Horn. Culberson Co., private property near Van Horn. 

Voshell. Cameron Co., Las Palomas Wildlife Management Area—Voshell Tract, Tx 
Farm road 511, 4 km SE of Brownsville. 

Warren. Tyler Co., US Hwy 69/287, small town. 

Welder Wildlife Foundation Refuge. San Patricio Co., US Hwy 77, 13 km NE of Sinton 
(19,768 ha). 

Westfield. Harris Co., small community off IH-45 N of Houston. 

Zapata. Zapata Co., private property nr. town. 


ACKNOWLEDGMENTS 


For certain citations and other useful information, I thank D. R. Davis, C. Dufay, R. 
W. Hodges, E. C. Knudson, E. G. Munroe, H. H. Neunzig, R. W. Poole, F. H. Rindge, 


VOLUME 41, NUMBER 4 Dok 


R. K. Robbins, and J. C. Shaffer. I am also grateful to May Elise Blanchard and Simone 
Blanchard Gérard for reviewing part of the manuscript. 


Roy O. KENDALL, 5598 Mt. McKinley Drive NE, San Antonio, Texas 78251-9609. 
Research Associate, Florida State Collection of Arthropods, Division of Plant Industry, 
Florida Department of Agriculture and Consumer Services, Gainesville, and Allyn 
Museum of Entomology of the Florida State Museum, Sarasota, Florida. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 238 


BOOK REVIEWS 


A REVISION OF THE GENUS HYPOCHRYSOPS C. & R. FELDER (LEPIDOPTERA: LYCAENIDAE). 
D. P. A. Sands. 1986. Entomonograph vol. 7. E. J. Brill/Scandinavian Science Press, 
Leiden. 116 pp., 2 color plates. About $38.25. 


Sands’s revision of the primarily Australian Region hairstreak genus Hypochrysops 
(Lycaenidae) is excellent. First and foremost, Sands knows these butterflies intimately. 
He has studied them in the field and museum, and clearly communicates his encyclopedic 
knowledge of them. There are 57 species—including 4 new ones—that can now be 
identified using the wing pattern keys to males and females. Almost half of the species 
are illustrated on two superb color plates. Sands figures male genitalia for each species, 
and comments on identification, larval foodplants, myrmecophily, and distribution. This 
information will be of interest to all students of Australian Region butterflies. 

Sands’s classification differs from that in previous books, such as D’Abrera (1977, But- 
terflies of the Australian Region, Lansdowne, Melbourne) because Hypochrysops had been 
confused with Waigeum. Hypochrysops species have red bands bordered with silver-green 
metallic markings while Waigeum resemble the distinctive “Danis wing pattern” shared 
by a variety of unrelated Australian Region blues and hairstreaks (Eliot, J. N. 1973. Bull. 
Brit. Mus. [Nat. Hist.] 28:399). Sands synonymizes these two genera because they do not 
differ structurally and because there are transitional forms. As this brief description 
indicates, Hypochrysops are beautiful butterflies. 

Two scientific highlights are noteworthy. First, Sands quantifies some of the variation 
that he finds in wing patterns and legs. The use of means and standard errors allows 
objective assessment of previous taxonomic results based on wing pattern and leg variation; 
hopefully Sands’s example will be followed by others. Second, Sands notes that “parts of 
the genitalia are almost membranous when specimens are freshly emerged and do not 
develop complete sclerotisation until about three days after emergence.” If this obser- 
vation, which I believe to be original, is widely true, it would account for some genital 
variation found in butterflies. 

Sands admirably discusses the systematic position of Hypochrysops among the hair- 
streaks. He presents an identification key to the four genera of the relatively homogeneous 
Hypochrysops Section. Clench (1955, Ann. Carnegie Mus. 33:261-274) considered these 
butterflies to be close relatives of the New World eumaeine hairstreaks, but Eliot (above) 
placed them in the tribe Luciini. Sands accepts Eliot’s classification, but I do not believe 
there is enough published information to resolve these differing views, one of the very 
few points on which I disagree with Sands. 

Despite my high praise, there are a few minor weaknesses with the book. Sands does 
not illustrate the female genitalia, an unnecessary omission. His classification is based 
mostly on overall similarity, not phylogeny, and I hope that he will consider a phylogenetic 
analysis as his next project. Along the same lines, he presents a diagnosis of Hypochrysops, 
but does not specify derived character states that “define” the genus. The biggest draw- 
back, however, is the $38.25 price tag for a slim 116 page book with two color plates. 
Apparently, it was priced for libraries, not individuals. However, those interested in the 
Lepidoptera of the Australian Region cannot afford not to have this excellent work. 


ROBERT K. ROBBINS, Department of Entomology, MRC NHB 127, Smithsonian In- 
stitution, Washington, D.C. 20560. 





Journal of the Lepidopterists’ Society 
41(4), 1987, 239 


NOCTUELLES ET GEOMETRES D EUROPE. DEUXIEME PARTIE. GEOMETRES. VOLUME IIJ— 
1917-1919. Jules Culot. Reprint edition, 1987. Apollo Books, Svendborg, Denmark. Order 
from: Apollo Books, Lundbyvej 36, DK-5700 Svendborg, Denmark. Vols. IJI-IV, DKK 
1380.00. 


This is an exact copy of the original volume, with 269 pp. and 37 color plates (figs. 1- 
771)—it is so identical, in fact, that the listings on the page of errat2 were not entered 
in the text of the reprint. The reprint edition is handsomely done, using glossy paper 
(which the original did not have) for both the text and plates; the latter are grouped 
together at the back instead of being randomly distributed throughout the text (as in the 
original). The figures were made from paintings and are all in color, always showing just 
half the moth, even though there is more than enough room to portray the entire insect; 
there has been a small loss of clarity and detail, and the colors do not always come up 
to the standards of the original, but the differences are usually small. 

This volume amounts to a descriptive and illustrated edition of Staudinger and Rebel 
(1901, Catalog der Lepidopteren des Palaearctischen Faunengebietes) as it pertains to 
Europe. The present volume does not have any descriptions or even indications of any 
higher categories, much less keys to separate them; the only descriptive material is on 
the specific level and below. No bibliographies or synonymies are given; you have to refer 
to Staudinger and Rebel for references to original descriptions and other literature. 

The Geometridae are covered in volumes 3 and 4 of this series. The family is arbitrarily 
chopped in half so that the two volumes will be approximately the same size. While this 
may make sense from a publisher's point of view, it doesn’t from a systematic one. Volume 
3 includes what today we call the Oenochrominae, Geometrinae, and part of the La- 
rentiinae (up to but not including Eupithecia). The Archiearinae is not included because 
Staudinger and Rebel placed Archieris Hiibner (Brephos auct.) as a separate family, the 
Brephidae. 

Unless you are fairly well acquainted with the European geometrids, this book poses 
a number of problems, beginning with the lack of definitions of the higher categories. 
Without generic descriptions or diagnoses it may be difficult to properly place a given 
species, as some members of this family look disconcertingly similar but may belong to 
entirely different genera.-If you are satisfied with the “picture book approach” you may 
be satisfied with Culot—but you take your chances on the identifications. 

Because the scientific names date from 1901, and no new scientific research has been 
included since then, this book may be of more interest to the antiquarian than to today’s 
collectors. I find a book published just before Culot’s to be much more useful; this is L. 
B. Prout’s Palearctic Geometridae (in A. Seitz, 1912-16 [English edition], The Macro- 
lepidoptera of the World, Vol. 4). Prout’s encyclopedic knowledge of the geometrids 
enabled him to do a better job of classification, and all the higher categories are defined. 
A modern treatment is given by W. Forster and T. A. Wohlfahrt in their Die Schmet- 
terlinge Mitteleuropas; these authors do not hesitate to illustrate venation, antennae, and 
the genitalia of both sexes whenever needed, as well as colored plates of the adults. 


FREDERICK H. RINDGE, Department of Entomology, American Museum of Natural 
History, New York, New York 10024. 


Journal of the Lepidopterists’ Society 
41(4), 1987, 240 


A HISTORY OF THE HOPE ENTOMOLOGICAL COLLECTIONS IN THE UNIVERSITY MUSEUM 
OXFORD WITH LISTS OF ARCHIVES AND COLLECTIONS. Audrey Z. Smith. 1986. Clarendon 
Press, Oxford. 172 pp. 


This short but informative book is divided into two parts. The first seven chapters tell 
the story of Rev. Frederick William Hope, his collections, and his quest to insure that 
they would be preserved, protected, and studied. The second and more significant part 
of the book contains five appendices. These list many of the holdings of the Hope 
Collections’ archives as well as names of individual donors and the collections they gave. 
Included in other appendices are a transcript of Rev. Hope’s Deed of Gift and an Oxford 
committee's reaction to the Deed. 

The author of this book is the Hope Librarian. Her attempt in the first third of the 
book to tell the history of Hope and his collections works fairly well. The text follows a 
logical sequence discussing Rev. Hope, curators of his collections, the collections them- 
selves, the Hope Professors, and a list of scholars associated with the Collections. Unfor- 
tunately, little detail is provided in the chapter examining Rev. Hope's life. As a wealthy 
gentleman-collector of the 19th century, his story may not differ from some others, but 
one cannot help wondering what his particular areas of interest were, and whether he 
made any lasting contributions to science through his activities. 

The other chapters also might have benefitted from additional detail, but they are 
sufficient to give the reader an idea of the importance of the Hope Collections. The 
chapter on the library of the Collections probably belongs in an appendix where other 
information regarding specific holdings has been placed. 

The major contribution of this book may be found in its first two appendices. For 
researchers interested in studying particular entomologists, Appendix A will be invaluable. 
Consisting of an alphabetical list of archival holdings, the list is not complete, but one 
glance at it indicates the vast quantity and quality of material in the Archives. Following 
each name in the appendix is a brief description of the material held. Holdings include 
correspondence, diaries, field notes, plates, paintings, and other media of entomologists 
from the late 18th to the late 20th centuries. Appendix B is an alphabetical list of donors 
with a description of the donation. It should be particularly valuable when used with 
Appendix A. Lepidopterists are common in both appendices. 

Lepidopterists will find A History of the Hope Entomological Collections to be a 
useful tool for research. Many archival repositories and museums contain priceless ma- 
terials that go unused because researchers are unaware of their existence. A publication 
such as this does researchers a true service by informing them of the holdings of a major 
facility. This is not a book that lepidopterists should be anxious to acquire for their 
personal libraries. It is, however, a generally well crafted work that all science libraries 
should own. 


TimoTHy D. Cary, Library, State University of New York at Stony Brook, Stony 
Brook, New York 11794-3323. 





Journal of the Lepidopterists’ Society 
41(4), 1987, 241-243 


INDEX TO VOLUME 41 


(New names in boldface) 


A History of the Hope Entomological Col- 
lections ..., book review, 240 

A Revision of the Genus Hypochrysops ..., 
book review, 238 

Acraea a. andromacha, 119 

Acraeinae, 119 

Acroceras zizanoides, 117 

Activity centers, 159 

Adult resources, 159 

Africa, 41 

Alkaloid, 166 

Anartia fatima, 75 

andromacha, a., Acraea, 119 

andromacha andromacha, Acraea, 119 

Announcement, 22 

Anolis, 141 

Apaturidae, 145 

appalachiensis, Bomolocha, 104 

Aquiioliaceae, 154 

arcas mylotes, Parides, 117 

Arctiidae, 166 

Aristolochia, 117 

Aristolochic acid, 141 

Asclepiadaceae, 13 

Asclepias fruticosa, 13 

Asteraceae, 23, 199 

Atalopedes nabokovi, 173 

Australia, 18, 23, 199 

Babcock, W. F., i 

Baccharis 
neglecta, 23, 199 
halimifolia, 23, 199 
sarathroides, 199 

Battus philenor, 141 

Behavior, 45, 116, 119 

Benrey, B., 141 

Biogeography, 29, 98 

Biological control, 23, 199 

Blanchard, A., obituary, 219 

Bomolocha appalachiensis, 104 

Boggs, C. L., 94 

Book reviews, 126, 128, 168, 169, 238, 239, 
240 

Boschniakine, 166 

Bowers, M. D., 75, 131 

British Columbia, 98 

Brown, R. L., 213 

Bucculatrix ivella, 23 

Buckett, J. S., obituary, 77 

Burns, J. M., 173 

Bush, M. B., 29 

Butler, L., 104 

Butterflies of Europe, Vol. 8, book review, 
128 


Calhoun, J. V., 1 
California, 121, 209 
California tortoise-shell, 121, 209 
California Butterflies, book review, 126 
californica, Nymphalis, 121, 209 
cardui, Vanessa, 116 
Cary, T. D., 240 
Catacolinae, 195, 212 
Catocala whitneyi, 212 
Cayman Islands, 145 
celtisana, Epinotia, 213 
clarkeata, Xanthorhoe, 98 
Colorado, 94 
Columbia, 65, 70 
confusa, Haploa, 166 
Conservation, 1, 29 
Corylaceae, 195 
Costa Rica, 75; 117 
Courtship, 119 
Crabtree, (R2C., 75 
Cyclopides paola, 41 
Danaidae, 13, 29 
Danaus plexippus, 13 
Defense gland, 187 
Dennis, R. L. H., 45 
Diatloff, G., 23 
digitalis, Penstemon, 166 
Distasteful, 141 
Distribution, 145, 195, 212 
Ecuador, 151 
Eggs, 131 
Eichlin, T. D., 154 
Endangered, 1 
England, 45 
Epinotia 
celtisana, 218 
vertumnana, 213 
Epstein, M. E., 119 
Eucosmini, 151, 213 
euryleon pleiades, Protesilaus, 70 
Eurytides thyastes, 114 
fatima, Anartia, 75 
Ferguson, D. C., 98 
Ferris, C. D., 128 
frutescens, Iva, 28 
fruticosa, Asclepias, 13 
garai, Gretchena, 151 
Genitalia, 173 
Geometridae, 98, 199 
Ghidiu, G. M., 154 
glaucus, Papilio, 83, 159 
Gonzalez, F. L., 145 
Graminae, 117 
Gretchena garai, 151 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Grey, P., Speyria collection, 123 Mexican Lepidoptera: Eurytelinae I, book 
Grossmueller, D. W., 159 review, 168 
halimifolia, Baccharis, 23, 199 Migration, 121, 209 
Hancock, D. L., 41 Miller, W. E., 22, 151 
Haploa confusa, 166 Mimicry, 141 
Hardesty, R. L., 116 Monarch, 13 
- Harrison, S. P., 75 mormonia, Speyeria, 94 
Heleantheae, 23 Multiple mating, 83 
Heliconiidae, 145 mylotes, arcas, Parides, 117 
Hemileuca lucina, 131 nabokovi, 
Henderson, R. M., 145 Atalopedes, 173 
Heraclides tasso, 108 Hesperia, 173 
Hesperia neglecta, Baccharis, 23, 199 
meskei, 173 neotropica, Juglans, 151 
nabokovi, 173 Neotropics, 114, 151 
Hesperiidae, 29, 41, 45, 145, 173, 187 nerva, Kedestes, 41 
Heterocampini, 187 New, T. R., 29 
Hibernation, 209 New Caledonia, 119 
hipparchus, Papilio, 65 Noctuelles et Géométres d Europe 
Hispaniola, 173 Vol. 2 (reprint), book review, 169 
Holloway, J., 169 Vol. 3 (reprint), book review, 239 
Hooper, J. D., 1 Noctuidae, 1, 104, 195, 212 
hydromeli, Litodonta, 187 Notodontidae, 187 
Iftner, D. C., 1 Nunez-Farfan, J., 141 
Ilex, 154 Nymphalidae, 1, 13, 29, 75, 94, 116, 119, 
Immature stages, 187 121, 145, 209 
Indonesia, 29 Nymphalis californica, 121, 209 
Infertility, 83 Obituaries, 77, 170, 219 
Introduction, 23 Ochlodes venata, 45 
Iva frutescens, 23 Odendaal, F. J., 141 
iwella, Bucculatrix, 23 Ohio, 1 
Java, 29 Olethreutinae, 151, 213 
Johnson, K., 65, 70, 108 Ontogeny, 187 
Juglandaceae, 151 Oviposition, 13, 117, 131, 159 
Juglans neotropica, 151 Palmer, W. A., 23, 199 
kathyae, Synanthedon, 154 paola, Cyclopides, 41 
Kedestes Papilio 
nerva, 41 glaucus, 83, 159 
protensa, 41 hipparchus, 65 
Kendall, R. O., 219 tasso, 108 7 
Knaus, R. M., 121 Papilionidae, 29, 65, 83, 108, 114, 117, 141, 
Lambremont, E. N., 121 159 
Larvae, 187, 195 Parides arcas mylotes, 117 
Lederhouse, R. C., 83, 159 Peacock, J. W., 1 
Leffler, S. R., 168 Penstemon digitalis, 166 
Leptocircini, 114 phaeocapna, Zale, 195 
Lesser Antilles, 145 phaon, Protesilaus, 65 
Lindroth, R. L., 166 philenor, Battus, 141 
Litodonta hydromeli, 187 Phylogeny, 214, 216 
lucina, Hemileuca, 131 Pieridae, 29, 145 
Lycaenidae, 1, 29, 145 Pipevine swallowtail, 141 
Lyonetiidae, 23 pleiades, euryleon, Protesilaus, 70 
Mating, 45, 119 plexippus, Danaus, 13 
Mattoni, R. H. T., 126 Polymorphism, 94 
Matusik, D., 65, 70, 108 Population structure, 13 
McCabe, T. L., 195 Predation, 75, 141 


Metzler, E. H., 1, 212 Prochoerodes truxaliata, 199 


VOLUME 41, NUMBER 4 


protensa, Kedestes, 41 
Protesilaus 

euryleon pleiades, 70 

phaon, 65 
Pupae, 114, 187 
Rausher, M. D., 141 
Relict population, 98 
Reviewers, manuscript, 124 
Rindge, F. H., 123, 239 
Robbins, R. K., 214, 238 
Roosting, 116 
Rosaceae, 131 
sarathroides, Baccharis, 199 
Saturniidae, 131 
Satyridae, 1, 29 
Schwartz, A., 145 
Scott, J.A., 216 
Scriber, J. M., 83 
Scrophulariaceae, 166 
Sesiidae, 154 
Sheppard, A. C., obituary, 170 
Shields, O., 209 
Shuey, J. A., 1 
Sobrevilla, C., 75 
Solomon, J. D., 154 
Spermatophore counts, 83 
Speyeria mormonia, 94 
Speyeria, P. Grey collection, 123 
Sphragis, 119 
Spiraea latifolia, 131 
Stamp, N. E., 131 
Stigma, 173 
Sudarman, H. K., 29 
Surveys, 1, 29 
Suzuki, Y., 13 2 


243 


Synanthedon kathyae, 154 
tasso 
Heraclides, 108 
Papilio, 108 
Taxonomy, 41, 65, 70, 98, 104, 108, 178, 
213 
Technical Comments, 22, 214, 216 
Territoriality, 45 
thyastes, Eurytides, 114 
Tiger swallowtail, 83, 159 
Tilden, J. W., 199 
Tortricidae, 151, 213 
Troidini, 117 
Vanessa cardui, 116 
Variation, 173 
Vasvary, L., 154 
venata, Ochlodes, 45 
vertumnana, Epinotia, 213 
Vickery, V. R., 170 
Watkins, R. A., 1 
Weed, 23, 199 
Weller, S. J., 187 
Wells, M., 75 
West, D. A., 114 
West Indies, 145 
whitneyi, Catocala, 212 
Williams, W. R., 45 
Wolfe, L. M., 75 
Xanthorhoe clarkeata, 98 
Young, A. M., 117 
Zale 
minerea, 195 
phaeocapna, 195 
Zalucki, M. P., 18 
zizanoides, Acroceras, 117 


Date of Issue (Vol. 41, No. 4): 5 January 1988 


‘ vith  atandannge A a 

oo mel 

+ S414 arrais et Ws i Ty ae 
wipe Aa ‘iN lena, ‘We . 1 ; af 





ve | I. gaum Pri nt Mime yey 


si. . watareh 

ei) sive? “nas 
Lhe et as 8 - 

vei | . (Late 


jas 


PL 
a 
~ 
~ 
it 
' i BS 
i 
‘ 
4 ioe + 
f ‘ 5 
\ 
rh 


> T 
Er ete Mel se cee a 


2 ea 


4 
: 
7 
t 
x 
4 
3 
x 









6 Smet pe %* oe ae 


Den Le a al PE < 


eae Pa oe Aa 


EDITORIAL STAFF OF THE JOURNAL 
WILLIAM E. MILLER, Editor 


Dept. of Entomology 
University of Minnesota 
St. Paul, Minnesota 55108 U.S.A. 


Associate Editors: 
M. DEANE BOWERS, BOYCE A. DRUMMOND III, DOUGLAS C. FERGUSON, 
ROBERT C. LEDERHOUSE, THEODORE D. SARGENT, ROBERT K. ROBBINS 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Categories are Articles, General Notes, Technical Comments, Book Reviews, 
and Obituaries. Short manuscripts concerning new state records, current events, and 
notices should be sent to the News, June Preston, Editor, 832 Sunset Drive, Lawrence, 
Kansas 66044 U.S.A. Journal contributors should prepare manuscripts according to the 
following instructions. 

Abstract: An informative abstract should precede the text of Articles. 

Key Words: Up to five key words or terms not in the title should accompany Articles, 
General Notes, and Technical Comments. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, entirely 
double-spaced, with wide margins, on one side only of white, letter-sized paper. Titles 
should be explicit, descriptive, and as short as possible. The first mention of a plant or 
animal in the text should include the full scientific name with author, and family. 
Measurements should be given in metric units; times in terms of the 24-hour clock (0930 h, 
not 9:30 AM). Underline only where italics are intended. 

Literature Cited: References in the text of Articles should be given as Sheppard (1959) 
or (Sheppard 1959, 1961a, 1961b) and listed alphabetically under the heading LITERATURE 
CITED, in the following format without underlining: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10:165-216. 


In General Notes and Technical Comments, references should be shortened and given 
entirely in the text as P. M. Sheppard (1961, Adv. Genet. 10:165-216) or (Sheppard, P. 
M., 1961, Sym. R. Entomol. Soc. London 1:238-30) without underlining. 

Illustrations: Only half of symmetrical objects such as adults with wings spread should 
be illustrated, unless whole illustration is crucial. Photographs and drawings should be 
mounted on stiff, white backing, arranged in the desired format, allowing (with particular 
regard to lettering) for reduction to fit a Journal page. Illustrations larger than letter-size 
are not acceptable and should be reduced photographically to that size or smaller. The 
author’s name and figure numbers as cited in the text should be printed on the back of 
each illustration. Figures, both line drawings and photographs, should be numbered con- 
secutively in Arabic numerals; “plate” should not be employed. Figure legends must be 
typewritten, double-spaced, on a separate sheet (not attached to illustrations), headed 
EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illustrations. 

Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material must be typed on separate sheets, and 
placed following the main text, with the approximate desired position indicated in the 
text. Vertical lines as well as vertical writing should be avoided. 

Voucher specimens: When appropriate, manuscripts must name a public repository 
where specimens documenting identity of organisms can be found. Kinds of reports that 
require vouchering include life histories, host associations, immature morphology, and 
experimental enquiries. 

Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author's changes at this time will be charged to 
authors at the rate of $2 per line. A purchase order for reprints will accompany proofs. 

Page charges: For authors supported by institutions, page charges are $20 per Journal 
page. For unsupported authors, page charges are $10 per Journal page with a $50 
maximum. Authors of Book Reviews and Obituaries are exempt from page charges. 
Authors unable to pay page charges for reasons such as difficulties with foreign exchange 
should apply to the editor for free publication. Four free publication slots are available 
annually in the Journal. 

Correspondence: Address all matters relating to the Journal to the editor. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A 





CONTENTS 


THE BIG SHIFT: NABOKOVI FROM ATALOPEDES TO HESPERIA (HES- 
PERIIDAE). john M. Burns ___.._._ 173 


LITODONTA HYDROMELI HARVEY (NOTODONTIDAE): DESCRIPTION 
OF LIFESTAGES. S, J. Weller: 187 


Hosts, BIOLOGY, AND DISTRIBUTION OF ZALE PHAEOCAPNA 
(NOCTUIDAE). . Tim L. McCabe ___... 195 


HOST SPECIFICITY AND BIOLOGY OF PROCHOERODES TRUXALIATA 
(GUENEE) (GEOMETRIDAE), A POTENTIAL BIOCONTROL AGENT 
FOR THE RANGELAND WEED BACCHARIS HALIMIFOLIA L. IN 
AUSTRALIA. W.A. Palmer & J. W. Tilden ae 199 


GENERAL NOTES 
Two related migrations of the California tortoise-shell butterfly in Mariposa 


County, California, in 1986.. Oakley Shields _____... Oa 209 
The type locality of Catocala whitneyi and reports of this species in Ohio. 

Eric H) Metzler uc re 212 
Correction of a name in the Epinotia vertumnana (Zeller) species-group 

(Tortricidae). Richard L. Brown 30 213 


TECHNICAL COMMENTS 
Logic and phylogeny: a critique of Scott’s phylogenies to the butterflies and 


Macrolepidoptera. Robert K. Robbins 2...) ee 214 

Logic and phylogeny: reply to R. K. Robbins. James A. Scott cece 216 
OBITUARY 

André Blanchard (1896-1986). Roy O. Kendall ...... 219 


Book REVIEWS 
A revision of the genus Hypochrysops C. ¢& R. Felder (Lepidoptera: Lycaeni- 


dae). Robert K. Robbins) 238 
Noctuelles et Géométres d'Europe. Deuxiéme partie. Géomeétres. Vol. I1I— 

1917-1919. Reprint edition. Frederick H. Rindge 2 ceccecceeeemeeee 239 
A history of the Hope Entomological Collections in the University Museum 

Oxford with lists of archives and collections. Timothy D. Cary ........ 240 


INDEX TO VOLUME 41 








ie " ae 
oo an 
~ 
} & Z 
} 1) 
; 
i 
; 
+ 
xX 
~ 
Wx ‘ 
wi 
y Y 
hd 
i 
fisy 
; 














a 
ts 
hy } 1 
; 
4 ' 
if 
f } 
j 
‘ 
) 
j 
Bey 
i 
ey, 
‘ 
ie 
( 
i 
j ; 
dul 
pit ae 
i r wo 
I wy 











esa 


“s 
Hint 


Pew 


Beas 
May 
See 

5! yy 

2s 


tal 
ad Pa 


b ny a 
Bd at hast 
Pagal oh ca 


rae 
ey 


% 
6 
* 


oF 


TWh tine 


yw 


it 
ne PAA 


. 


4: 


at 
4 
yay 
Bois 
ry es 
oe 

ue 
ew 


£ 
yearay 


a's 
’ 


* 
‘ 
i 


Are NA 
Pee 
GY Pay, 
Ae ole 
» 
442 
YY Owe aya” 


Boa 
osu 
cae 
MA 
bewoy 
ane 


15 ly